Bulletin of the Museum of Comparative Zoology HARVARD UNIVERSITY Vol. 133, No. 10 THE AMEIVA (LACERTILIA, TEIIDAE) OF HISPANIOLA. II. GEOGRAPHIC VARIATION IN AMEIVA CHRYSOLAEMA COPE By Albert Schwartz and Ronald F. Klinikowski CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM March, 1966 OCR text unavailable for this page.Bull. Mus. Comp. Zool., Harvard Univ., 133(10): 425-487, March, 1966. No. 10 — The Ameiva {Lacertilia, Teiidae) of Hispaniola II. Geographic Variation in Ameiva Chrysolaema Cope By Albert Schwartz 10,000 SW 84th Street, Miami, Florida 33143 AND Ronald F. Klinikowski 127 Spring St., Reading, Pennsylvania INTRODUCTION The largest of the three species of Hispaniolan ground lizards, Ameiva chrysolaema Cope, was described in 1868 with the type locality ''Gonave Island." Three other names were then ap- plied to this species in rapid succession: vittipunctata Cope 1871, affinis Fischer 1883, and regularis Fischer 1888. Of these, vittipunctata, as Cochran (1941: 275-276) has made clear, was apparently described by Cope from a young specimen (snout- vent length 88 mm) which was part of the same series from which he himself had taken the type of chrysolaema. Cope, how- ever, gave a different type locality — ' ' city of Santo Domingo ' ' — for vittipunctataA In her revision of the species in ' ' Herpetology of Hispaniola, ' ' Cochran (1941: 275-292) considered vittipunctata Cope a strict synonym of A. chrysolaema Cope. A. regularis Fischer was re- garded as a strict synonym of affinis Fischer, which was accepted as a valid mainland subspecies (a new status since affinis had previously been regarded either as a full species or a synonym). Prior to 1941 two subspecies of A. chrysolaema had been de- scribed from satellite islands of Hispaniola : A. c. woodi Coch- ran from He de la Tortue and A. c. ahhotti Noble from Isla Be- ata. These Cochran considered recognizable. A third subspecies, A. c. hoekeri Mertens (1938: 338), however, from the mainland at Fondo Negro, Republica Dominicana, was rejected as a syno- nym of A. taeniura. 1 Cochran (1941 : 245) noted that the type locality of Cele^tus f= Diploglos- SU8) weinlandi, also described by Cope and said by him to have been collected on He de la (ionave, was incorrect and the s|ipcimen actnallv came from the mainland within 25 miles of Port-au-Prince, Haiti. Since A. C. Younglove who collected the type of C. tveinlanrti in 1868 also collecred the type of A. rhrysn- laema, it is appropriate, as Dr. Cochran has done, to restrict the type locality of A. chryftolaema to "within 2.t miles of Pnrt-aii-I'rince." If the type of A. vittipunctata is actually part of the same series as the type of A. chrysolaema, the type locality of the former should likewise be considered the same as that of the latter, despite Cope's statement that it came from the city of Santo Domingo. 428 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY The discussion below departs radically from Cochran's revi- sion in rejectinpr affinis while recoonizing" many other mainland subspecies. In large part this has been due to the much greater amount of material available to us. Six hundred and fifty-five carefully documented and noted specimens of A. chrysolaema have been collected by ourselves and associates betvreen June 1962 and September 1964. Large fresh series of this lizard are now availal)le to us from southern and central Haiti, from He de la Tortue and Gonave, and from the whole of its range in the Republica Dominicana. These speci- mens are in the Albert Schwartz Field Series (ASFS) and the collection of Richard Thomas (RT). They have been amassed through the efforts of ]\Iiss Patricia A. Heinlein, and Messrs. David C. Leber and Richard Thomas. All deserve our commen- dations and thanks, most especially Messrs. Leber and Thomas who made special efforts on our liehalf to secure topotypes of A. c. ahboffi on Lsla Beata. Li addition, we have examined 417 specimens in the following collections : American ^Museum of Natural History (AMXH), Carnegie ]Museum (C^I), ]\Iuseum of Comparative Zoology (MCZ), Museum of Zoology, University of Michigan (U]\OIZ), and United States National Museum (USNM). To the curators and their assistants — Charles M. Bogert and Miss Grace M. Tilger, Neil D. Richmond, Ernest E. Williams, Charles F. Walker and George R. Zug, Doris M. Coch- ran and James A. Peters — we wish to express our appreciation for allowing us to examine pertinent specimens under their care. Paratypes of new forms have been deposited in the Museum of Natural History, University of Kansas (KU), and the University of Illinois Museum of Natural History (UIMNH), as well as in the above collections. The Harvard collections once again have been indispensable ; through the efforts of Dr. "Williams, large and well preserved series from northern Haiti have been made available to us; without these our interpretation of the north- western Haitian situation would have been not merely difficult, but rather impossible. In the matter of literature, Edmond V. Malnate has been most helpful and we are grateful for his co- operation. The illustrations are the work of the junior author. SYSTEMATIC ACCOUNT The Species as a Whole Amciva clirysolacma may be defined as follows: 1) a large species of the genus Amcii'a with snout-vent length to 160 mm in males and 134 in females; 2) dorsal caudal scales keeled and SCHWARTZ AND KLINIKOWSKI : AMEIVA 429 straig-lit; 3) ventrals in 10, 11, or 12 transverse rows and in 33 to 41 long-itndinal rows; 4) fourth toe subdipital seales from 66 to 101; 5) femoral pores 24 to 52; 6) fifteenth eandal verticil with 30 to 52 scales; 7) dorsal pattern consistino- of one of the following: a) a series of dorsal yellow to huffy longitudinal lines on a brown, tan, grayish tan, greenish, or lilackish ground color, the lines (straight or wavy) at times modified into dashes, dots, or fused with one another to give ultimately a median dor- sal longitudinal band, b) a rather uniform covering of brightly colored (blue, orange, yellow) spots on a dark background, c) a pale ground with dark vermiculations and tigroid vertical lateral bars, or d) completely or almost unicolor dorsally without any striking jiattern elements; and 8) hemipenis extending to about the seventh to ninth caudal verticil, sulcate surface naked, sulcus bifurcating apically, the branches ending in two poorly defined scalloped apical discs, non-sulcate surface entirely flounced, the flounces extending to the margins of the sulcate sur- face, a small smooth triangular area on the non-sulcat(> side which divides the flounces for about one-third the length of the organ into two fields of flounces which correspond to the apical discs. The center of the distriliution of A. chrysolacma is in the (Jul de Sac-Valle de Xeiba region of llispaniola; the species is rep- resented by large series and from numerous localities in this gen- eral region. From here, A. cJirysoIacDia extends westward to the vicinity of Leogane on the Tiburon Peninsula, and eastward as far as San Pedro de Macoris in the Pepublica Dominicana. From Leogane to San Pedro de ]\lacoris, there is a set of localities which implies a more or less continuous distribution. From this basic center, populations extend northAvestwarcl along the shore of the Golfe de la Gonave into the valley of the Riviere de I'Arti- lionite, and in the central portion of Haiti at least as far north as the vicinity of Mirebalais. In the Kepublica Dominicana, there is practical continuity of the main southern mass of the species north of the Sierra de Xeiba in the Valle de San Juan and thence to the Dominico-Haitian 1, order near Pedro Santana and immediately across the border at Cerca-la-Source in Haiti. A second major center lies in the northwestern portion of Haiti and extends thence eastward into the Valle de Cibao in the Republica Dominicana. Whether these two major populations are com- pletely isolated from one another is unknoAvn ; there is at least no contact in the Republica Dominicana, since the Cordillera Central stands between the two main regions, and the central valley along the eastern slopes of this range is unoccupied by chrysolaema. 430 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY There are several apparently disjunct populations in southern Haiti and the Republiea Dominicaua ; in the former country there is a single specimen from Aquin to the west along the Tiburon Peninsula, far removed from the nearest records from Leogane. In the Republiea Dominicana there are specimens from Juanillo near the eastern extremity of the island. Although there are no records from between San Pedro de Macoris and Juanillo, the species has presumably occurred in that region fairly re- cently (and may still occur as isolated populations), since it oc- curs on Isla Catalina and Isla Saona. It is at present unknown from the adjacent mainland in each case. There were two speci- mens reported (Cochran, 1941:282) from Ile-a-Vache but these cannot now be located ; although in two extended visits to this island we secured only the very abundant A. tacniura Cope, it is not altogether impossible that A. chrysolaema occurs there as well. Finally, there are populations south of the Sierra de Bao- ruco-Massif de la Selle which are completely cut off by these ranges from their more northern relatives ; this same phenomenon has now been noted in several other species of reptiles from this area. These mountains, as well as the virtually non-existent coastal plain along the eastern shore of the Peninsula de Bara- hona, form an inescapable trap for several reptiles in the south- ern portion of the Peninsula. For the remaining satellite islands, A. chrysolaema is repre- sented by abundant material from He de la Gonave and occurs also on He de la Tortue, Isla Beata, and the Siete Hermanos islands off the northwestern portion of the Republiea Domini- cana. Of the three species of Hispaniolan Anieiva, none has so broad a range as does A. chrysolaema. Nonetheless, there is geo- graphic evidence that even this species is retracting its range ; the apparent absence of chrysolaema from much of the south- eastern portion of the Republiea Dominicana, but its occurrence on off-shore islands in this region, suggests a formerly more widespread distribution in this area. A. chrysolaema is absent from the distal third of the Tiburon Peninsula in Haiti, and apparently from much of the central i^or- tion of that country as well. In the Republiea Dominicana it does not occur in the Cordillera Central nor the Sierra de Neiba, and is absent from the central and eastern portions of the country- except along the southeastern coast. Interestingly, despite its occurrence in the Valle de Cibao, it does not occur along the northcentral coast of the Republiea Dominicana. The Cordillera SCHWARTZ AND KLINIKOWSKI : AMEIVA 431 Septentrional acts as an effective barrier in this region. Much of the eastern Republica Dominicana is mesic (in fact, the area of highest rainfall in the country occurs in this region), and since A. chrysolaema is distinctly a lizard of xeric habitats, this one factor may well have prevented its expansion into this region and onto the Peninsula de Samana. However, its absence along the coast from Cabo Engaiio westward is strange, since this coast is arid and appears suitable for these lizards. Considering the disjunct nature of the populations of the species in extreme east- ern Hispaniola, it is possible that it never occurred in this region or that it has already retreated from this suitable coastal area. A. chrysolaema lives in xeric regions. It is abundant in the Cul-de-Sac plain below sea level and on the Peninsula de Bara- hona. It occurs also in the dry Valle de San Juan, at elevations of about 1000 feet. Although more tolerant of less xeric situa- tions than A. lineolaia, the two often occur together, with A. Uncolata inhibiting more open, cactus-studded, sandy regions, and A. chrysolaema preferring slightly more shady areas, such as adjacent copses or thickets of Acacia. If lowland woods are present, A. chrysolaema may invade them; the woods may not be dense nor with abundant ground cover. Maritime deciduous for- ests along the mangrove border (but usually not the mangroves themselves) offer a suitable habitat. Scrub-lands and open beaches with some cover are often adetiuate. In its relationships with the other two species, A. chrysolaema most often occurs with A. lineolata as noted above. On occasion, however, A. chrysola- ema occurs with the shade-loving A. taeniura. In such instances, chrysolaema appears to be tlie secondary invader of a habitat which is the preferred habitat of taeniura; in one such case near Oviedo on the Peninsula de Barahona, taeniura kept strictly to the open dry forest, whereas chrysolaema occurred almost exclu- sively along the edges of the woods where they abutted on a dry mangrove flat. The latter is likely the more preferred habitat of chrysolaema, but during the heat of the day this species was not averse to foraging in marginal forested situations. Characters studied We have examined a total of 1072 specimens of A. chryso- laema (in contrast to 42 examined by Barbour and Noble, 1915, and 198 by Cochran, 1941) ; of these, 655 are specimens collected by ourselves and parties at various times, and upon which we have extensive data on coloration and pattern. Of the races 432 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY recognized and described in the present paper, we have seen liv- ing or freshly killed specimens of all but two (the race from northwestern Haiti, and the race from extreme eastern Repvib- lica Dominicana). We feel that data on coloration and pattern are absolutely indispensable for any modern worker on the genus Ameiva. Old, faded, or (even worse) badly discolored specimens are completely useless except for scale counts and measure- ments, and analyses of populations must rest heavily and se- curely upon data from living or freshly killed animals. We have taken counts of rows of longitudinal and transverse ventrals, fourth toe subdigital scales, femoral pores, and scales in the fifteenth tail verticil (see Tables 1-4). Of these, only the numlier of transverse rows of ventrals can be used (partially) to characterize subspecies — i.e., having either 10 or 12 transverse rows of ventrals. Xo ]K)pulation has all specimens with 10 or all with 12 rows. However, there is most often a preponderance of one or the othei- in any particular sample, and we have used this modal number as typical of the race in question (Table 2), un- less the sample is rather small or the two categories differ by only a very few individuals. Of least value systematically is the number of longitudinal rows of ventrals. In the entire lot of A. chrysolacma examined, this figure varies from 33 to 41. The means for the 15 popula- tions described herein vary from 38.3 to 36.5. Data for longitu- dinal rows are presented in each case, but these data are not em- phasized. The counts of fourth toe scales, femoral pores, and fifteenth verticil scales show some trends, although in almost all cases the amount of overlap is rather large between most populations. Dif- ferences of means, however, between those races which rank first and last in each category may be rather striking (see Tables 2. 3 and 4). The largest difference between the highest and lowest populations is in fourth toe scales, where the high population has a mean of 91.0, and the low 77.8 — a difference of 13.2 scales. For facility Ave have combined in all cases the fourth toe scales from both feet and the femoral pore counts for both legs into one figure for each specimen ; we do not feel that this weakens the use of the data and it may well intensify slight differences, which would otherwise be almost unnoticeable. We have given the means and extremes for these three scale counts for each subspecies ; the differences, if any, obviously are mean differ- ences, since overlap of ranges is great in most cases. schwartz and klinikowski : ameiva 433 The Recognizable Subspecies Ameiva chrysolaema chrysolaema Cope, 1868 Ameiva chrysolaema Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 20:127 (type locality — "He de la Gonave" = within 25 miles of Port-au- Prince, fide Cochran, 1941:275). Ameiva vittipunctata Cope, 1871, Proc. Acad. Nat. Sci. Philadelphia, 22:220 (type locality — "city of Santo Domingo" = within 25 miles of Port-au-Prince; see Cochran, 1941:275-76 for discussion of rationale for this restriction). Ameiva affinis Fischer, 1883, Beschreibungen neuer Eeptilien, [Separat- Abdruk aus dem] Osterprogramm des akademischen Gymnasiums, Ham- burg, p. 1 (type locality — "Haiti"). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of very large size (males to 160 mm, females to 130 mm snout-vent length), usually 12 transverse rows of ventrals, moderate number of fourth toe subdigital scales, high number of femoral pores and of scales in the fifteenth tail verticil ; dorsal pattern consisting of about six longitudinal yellow lines and/or yellow dots arranged in series (Fig. 1, left), and a black gular band which may involve the chest and undersides of the arms. Distribution: From St. Marc (and including the "Artibonite Valley") on the north, southeast along the shore of the Golfe de la Gonave, east throughout the Cul-de-Sac plain to the environs of Etang Saumatre (Manneville, Ganthier, Fond Parisien), and west on the Tiburon Peninsula as far as the vicinity of Leogane (Pere) ; an isolated specimen from Aquin, Dept. du Sud, Haiti (Fig. 11). Discussion : A. c. chrysolaema is distinctly the largest and most bulky of the races of the species. Males reach a snout-vent length of 160 mm and females 130 mm. Color notes on a series from Eaux Gaillees in the Haitian Cul de Sac show the situation as far as coloration and pattern are concerned. Males from Eaux Gail- lees were noted as dark brown to reddish brown dorsally, espe- cially reddish on the shoulders and head (which may also be grayish). Lores and cheeks with gray blotches. The back has either a series of six yellow lines and yellow dots in the inter- spaces between the lines, or has six rows of lemon yellow spots. The lateral fields are black with or without a longitudinal series of yellow spots. The lower sides have large yellow spots as well, and the lateralmost belly plates are blue-spotted. The ventral ground color is dull blue-gray, the throat pale orange (Maerz and Paul, 1950, PL 9D7). There is a black gular band, 434 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Fig. 1. Left, Ameiva chrysolaema chrysolaema, ASFS X2162, 3.9 mi. NW Ganthier, Dept. de 1 'Quest, Haiti. Right, Ameiva c. umiratilis, holo- type, MCZ 77231, Baraliona, Barahona Prov., Eepublica Dominicana. which may expand posteriorly to cover the chest and under- sides of the arms. Some adult males have the shoulders and neck blackened, so that in these regions the yellow lines are much dulled and obscured. The females are like the males except that the lines are yellow and prominent anteriorly. The venter is dull blue-gray. The black gular collar is present but less pro- nounced than that of the males, and the throat is orange but paler than that of males. SCHWARTZ AND KLINIKOWSKI : AMEIVA 435 A series from Diquiiii to the ^vest of Port-au-Prince was col- ored much as the Eaiix Gaillees material. The dorsal ground color in males was brown with yellow dots arranged in lines or with yellow lines additionally present. There is a black lateral field with yellow spots. The sides of the head and axillae had vivid and prominent blue to blue-green blotches. The heads were dull reddish brown to dull orange, with orange-pink throats. The ventral ground color was grayish to orange with blue spots on the sides of the abdomen. The hindlimbs were dotted with yellow, the forelimbs with blue-green. Females resemble the males, but the dorsal lines or dots are less bright and prominent. From the above descriptions it is obvious that, despite some diiferences in details, these two populations (as well as many others throughout the range of chrysolaema) share a communitj' of dorsal markings — the longitudinal series of yellow^ lines and/or dots. Cochran (1941: pi. 8E) showed a dorsal view of the type of A. chrysolaema. The six dorsal lines, in this case partially fragmented into series of longitudinal dashes, are quite distinct. There is no doubt that the type of A. chrysolaema did indeed originate in the vicinity of Port-au-Prince, since only this subspecies occurs anywhere near that city. Specimens from He de la Gonave are much duller, less prominently marked, lack fragmentation of the dorsal lines, and are not referable to the nominate form. The longitudinal ventrals vary between 35 and 40 (mean 37.7) and these scales are most often arranged in 12 transverse rows (67.9 per cent), with 29.2 per cent having 10 transverse rows and 2.9 per cent having 11 rows. The variation in number of transverse rows depends primarily on whether the lateralmost of the enlarged ventral scales is sufficiently large to be con- sidered a ventral; we have so considered it if its length (longi- tudinally) is equal to that of the next inner adjacent row, and if its width (transversely) is equal to at least half that of the next inner adjacent row. Occasional specimens may also have one or two rows of the belly plates divided, thus attaining counts of 11 or 12 in another fashion. The fourth toe subdigital scales range from 76 to 101 (mean 86.7) and the femoral pores range from 33 to 50 (mean 43.7). The scales in the fifteenth caudal verticil vary between 37 and 51 (mean 44.4). Within the range ascribed to A. c. chrysolaema, there are vari- ous relatively minor pattern variants which we consider as 436 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY part of the normal variation of the subspecies. Specimens from St. Marc at the northwesternmost extreme of the range are very boldly lined longitudinally; the same is true of two indi- viduals from Manneville. In both cases there are specimens from adjacent localities or from the same locality which have more typical chrysolaema markings. The single individual from Aquin (USNM 72614) is unique in several (possibly significant) features. The longitudinal lines are composed of dots which also have a more or less transverse arrangement, so that the back has a rather conspicuous trans- versely banded appearance, a condition seen in no other A. c. chrysolaema. The black lateral field is obscured, and there is no black gular band, the entire throat, chest and undersides of the arms being unmarked. The specimen is an adult male with a snout-vent length of 146 mm, ventrals in 38 longitudinal and 12 transverse rows, 95 subdigital fourth toe scales, 41 femoral pores, and 43 scales in the fifteenth caudal verticil. None of these counts will distinguish the specimen from A. c. chrysolaema, although we attach no particular significance to this fact. Considering the wide gap between the known localities of A. chrysolaema between Pere near Leogane and Aquin (a distance of some 90 kilometers) and the fact that the Aquin specimen comes from the southern, in contrast to the northern, side of the Tiburon Peninsula, it is likely that this single individual comes from a population which is distinct from A. c. chrysolaema. AVithout additional material, and especially lacking careful data on col- oration and pattern in life, we are unwilling to name this single Aquin specimen as distinct from A. c. chrysolaema. The character ascribed by Cochran (1941:277) to differenti- ate A. c. affinis — i.e., the interparietal being larger than the adjacent scales — we find to be completely untenable. Dr. Coch- ran has also shown (1941:291) that the scale counts of affinis fall within the known range of A. c. chrysolaema. Specimens which she assigned to affinis were reported (1941:292) from Momance, Manneville, and Pere in Haiti. Other specimens (p. 282) from Manneville were assigned to c. chrysolaema. In a series of twenty-seven specimens from Fond Parisien and the eastern Cul de Sac, for example, six have the interparietal larger than adjacent scales and the balance have the interparietal smaller. The same situation applies to series from other locali- ties within the range of A. c. chrysolaema, and if we accept affinis as differentiated by this character alone, then the races SCHWARTZ AND KLINIKOWSKI : AMEIVA 437 chrysolaema and affinis are broadly and randomly sympatric. No features of pattern or coloration will distinguish specimens with smaller interparietals from those with larger interparietals, and we cannot detect any other constant scale feature which will distinguish two forms in this region. For this reason we con- sider affinis Fischer as a synonym of chrysolaema Cope. The reasons for considering vittipunctata Cope as a synonym of chrysolaema have been outlined in the introduction. Specimens examined: Haiti, Dept. de I'Artihonite, "Artibonite Valley" (not mapped), 1 (USNM 75921) ; St. Marc, 5 (USNM 59079, MCZ 58012-13, 65351, AMNH 49766) ; Bept. de VOuest, 2.2 mi. (3.5 km) SW Trou Forban, 1 (ASFS X1927) ; between Arcahaie and Trou Forban, 1 (MCZ 51433) ; 6.3 mi. (10.1 km) NE Arcahaie, 3 (ASFS X1928, X1930, X1946) ; 13 mi. (20.8 km) SW Arcahaie, 1 (ASFS X1938) ; Port-au-Prince, 12 (AMNH 49637-38, MCZ 13839, 59495-502, 69420) ; Carrefour Feuille, Port-au-Prince (not mapped), 1 (MCZ 65810) ; Delmas, 2 (MCZ 65808-09) ; Petionville, 1 (USNM 59078) ; 10 mi. (16 km) SW Port-au-Prince, 1 (UMMZ 92197) ; 3.5 mi. (5.6 km) E Croix des Bouquets, 15 (ASFS X2197-211) ; Eaux Gaillees, 33 (ASFS X1651-83) ; Manneville, 11 (MCZ 8621-23, 8625, 8629-33, 8614, 8618) ; 3.9 mi. NW Ganthier, 18 (ASFS X2153-70) ; 1.3 mi. (2.1 km) NW Fond Parisien, 3 (ASFS X2174-76) ; 0.4 mi. (0.6 km) SE Fond Parisien, 7 (ASFS X2189-95) ; Hatte Latham (not mapped), 1 (MCZ 51424); Diquini, 36 (ASFS X2381-407, MCZ 6292, 8706, 8649-51, 8653-54, 8658-59) ; Momance, 8 (MCZ 8634-35, 8638-41, 8649, 20875) ; Qa Ira, 2 (MCZ 64919-20) ; Pere, 3 (MCZ 13271-73) ; Dept. du Sud, Aquin, 1 (USNM 72614). Ameiva chrysolaema umbratilis,^ new subspecies Ilolotype: MCZ 77231, a subadult female, from Barahona, Barahona Province, Republica Dominicana, taken 25 July 1963, one of a series taken by native collectors. Original number X9721. Paratypes: All from the Republica Dominicana, as follows: MCZ 81000-04, USNM 152558-60, KU 79861-64, UIMNH 56886- 89, RT 738-39, same data as holotype; ASFS X9568-69, Bara- hona, Barahona Prov., 24 July 1963, native collector; AMNH 37943-49, 38133-39, Barahona, iBarahona Prov., 12 October 1922, 1 From the Latin for "remaining in the shade." 438 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY G. K. Noble; AMNH 63191, 63193, Barahona, Barahona Prov- ince, 10-19 July 1932, W. G. Hassler; MCZ 58019, Barahona, Barahona Prov., 13 July 1932, W. G. Hassler; MCZ 43813-14, Barahona, Barahona Prov., 18 July 1932, W. G. Hassler ; ASFS V199-200, 4 km NW, 1 km SW Barahona, Barahona Prov., 5 August 1963, A. Schwartz, R. Thomas; AMNH 49837-38, "Palo- mino Spring's, nr. Barahona" (not mapped), Barahona Prov., 15 August 1935, W. G. Hassler. Associated specimens : Rcpuhlica Dominicana: Independencia Prov., 6.5 mi. (10.5 km) NE Jimani, 1 (ASPS X9507) ; 4.4 mi. (7.0 km) SE Jimani, 3 (ASFS X9515-17) ; 13 km SW La Descubierta, 3 (ASPS X9364-66) ; 5 km E La Descu- bierta, 5 (ASFS X9354-58) ; Las Baitoas, 1 (MCZ 58776) ; 22 km SE Duverge, 7 (ASFS X9928-34) ; 1 km W El Naranjo, 1000 feet, 2 (ASFS X9943-44) ; northv^^est side, Laguna del Rincon, 1 (MCZ 58779) ; Guayabal, 9 km N Postrer Rio, 2 (MCZ 57732- 33) ; Baoruco Prov., Jaragua, 5 (ASFS X9469-72, RT 713) ; 0.7 mi. (1.2 km) E El Estero, 2 (ASFS X9467-68) ; 0.8 mi. (1.3 km) SW Neiba, 15 (ASFS V252-64, RT 775-76). Diagnosis: A subspecies of A. chrysolacma characterized by a combination of moderate size (males to 130 mm, females to 112 mm snout-vent length), usually 10 (but often 12) transverse rows of ventrals, moderate number of fourth toe subdigital scales, low number of femoral pores, and high number of scales in the fifteenth caudal verticil ; dorsal pattern consisting of dull grayish brown to greenish black dorsal ground color with a series of eight to ten dorsal longitudinal lines composed of small and numerous dull yellowish to tan dots (Fig. 1, right), and a black gular band which in adult males may involve the chest and undersides of the arms. Distribution : The Valle de Neiba from just east of Jimani to the vicinity of tlie city of Barahona, Republica Dominicana (Fig. 11).' Description of type: A subadult female with the following measurements and counts : snout-vent length 93 mm, tail 184 mm ; ventrals in 37 longitudinal and 10 transverse rows ; fourth toe subdigital scales 39 and 41 (total 80) ; femoral pores 18 and 17 (total 35) ; 43 scales in the fifteenth caudal verticil. Dorsal ground color grayish brown in life, head gray, shoulders greenish ; ten rows of dull yellowish dorsal dots, the dots in each series virtually confluent with one another, giving a vague wavy line ; lateral fields obscure darker gray with scattered buffy dots. SCHWARTZ AND KLINIKOWSKI : AMEIVA 439 Throat pale purplish orange, followed by a black gular band which extends slightl}' onto the chest and onto the underside of the f orelimbs ; ventral ground color grayish blue ; tail dull grayish brown above and grayish blue below, with some scattered darker scales dorsally. Variation: See tables. The characters of umhratilis are best expressed in populations from the eastern section of the Valle de Neiba ; however, even the most western specimens from the vicin- ity of Jimani are in no way comparable to Haitian A. c. chryso- laenia. The dorsal ground color was noted in life as being green- ish black (Jimani, La Descubierta, Jaragua), greenish brown (Duverge, El Naranjo), brown (El Estero), and grayish brown (Barahona). The dots in the dorsal longitudinal lines are tiny and very often confluent, giving almost a vermiculate appearance to the dorsal band; the dots vary in color from greenish (Jimani), creamy (La Descubierta), pale yellow to pale green (Duverge), yellow (El Estero), grayish yellow (Jaragua), or dull yellowish to tan (Barahona). The lateral fields are usually dull and inconspicuous, hardly darker than the lateral coloration ; they often include a row of yellow to creamy spots and are not outlined either above or below by pale and prominent longitudinal lines. The ventral coloration varies from pinkish gray and grayish orange to grayish blue, with specimens having the brighter colors known from the western extremity of the range. The throats are dull pinkish gray and grayish orange to dull purplish or dull orange. The l)lack gular band is invariably present and may, in adult males, expand to cover much of the chest and anterior abdomen and underside of the forelimbs. The upper surfaces of the limbs are usually unspotted, but if there are a few scattered dots these are blue on the forelimbs and yellow on the hindlimbs. Comparisons: The coloration and pattern of chrysolaema and xmhratilis are strikingly different; even in the western portion of the range of umhratilis, no specimen approaches closely the vivid dorsal coloration and pattern of the nominate race. The extreme condition in the eastern Valle de Xeiba contrasts strongly with the condition at Fond Parisien, for instance, and specimens from Jimani and La Descubierta are much more like individuals from Barahona in having fine dorsal dotting and generally more drab colors than they are to specimens from Fond Parisien. Umhratilis is a smaller lizard; no specimen of cither sex of this race achieves the much bulkier and larger 440 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY size of chrysolaema. This is certainly not a sample artifact since large series of both forms are at hand. The tendency for vmhra- tilis to have 10 versus 12 transverse rows of ventrals, as in chrysolaema, is of interest, although in umhratilis the specimens are almost equally divided between the 10- and 12-row condi- tions. In number of longitudinal ventral rows, the two races are comparable {chrysolaema 37.7, umhratilis 37.1). In number of femoral pores these two races differ strongly, with a mean of 43.7 in chrysolaema and 35.6 in umhratilis. Chrysolaema aver- ages slightly higher in counts of femoral pores and fifteenth verticil scales. Remarks: The occurrence of two very distinct races of A. chrysolaema in the Cul de Sac-Valle de Neiba complex is sur- prising. Aside from the more mesic eastern and western ends of this long xeric valley, the conditions throughout are quite com- parably severe. Interestingly, the Valle de Neiba is greatly con- stricted just to the east of Jimani ; it is possible that this narrow neck (7-10 km) has been effective in separating these two races. Specimens from the Republica Dominicana to the northwest of Jimani may well be assignable to A. c. chrysolaema. Ameiva chrysolaema boekeri Mertens, 1938 Amciva chrysolaema hoeTceri Mertens, 1938, Senckenbergiana, 20:338 (type locality — south of Foiido Negro, lower Rio Yaque del Sur, Bara- hona Province, Republica Dominicana). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of moderate size (males to 126 mm, females to 111 mm snout-vent length), usually 10 transverse rows of ven- trals, moderate number of fourth toe subdigital scales, low num- ber of femoral pores, and high number of scales in the fifteenth verticil ; dorsal pattern of two phases : ( 1 ) back yellowish brown, grayish tan, to olive, and without pattern and often without any indication of lateral fields, or (2) dorsum colored as above but with faint paler marblings or longitudinal lines and a fairly prominent black to dark gray lateral field (Fig. 2), and a black gular band which may involve the chest and underside of the arms. Distribution: North of the Rio Yaque del Sur in extreme eastern Valle de Neiba, north and east to north of Azua and east to the vicinity of Bani, in the Llanos de Azua, Republica Dominicana ; intergrades with the next subspecies to the north- west in the vicinity of Hato Nuevo, Azua Province (Fig. 11). SCHWARTZ AND KLINIKOWSKI : AMEIVA 441 Fig. 2. Left, Ameiva c. hoekeri, ASFS X7811, 10 mi. NW Bani, Peravia Prov., Eepiibliea Dominicana. Eight, Ameiva c. boeTceri, ASFS V689, 15.2 mi. S San Jose de Ocoa, Peravia Prov., Eepubliea Dominicana. Discussion: A. c. hoekeri was described on the basis of four- teen lizards from Fondo Negro. Of these, seven males were dorsall}^ patternless (including the type), four males showed a " chriisolaema"-\ike pattern, and the final male was considered by Mertens (1939:72) to resemble the Beata race ahhotti — i.e., it was dorsally dotted. The two paratypic females were " chryso- toema "-like as well. We have examined a single paratype of hoekeri (MCZ 44757) and eighty-six other specimens from the range ascribed by us to hoekeri above. Of ten localities, only 442 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY two have "pure" or almost "pure" hoekeri populations (i.e., patternless), viz., a series of five specimens from the west side of Punta Martin Garcia, Barahoua Province, and a series of twenty-one from 10 mi. XW Bani, Peravia Province. Additional specimens (four) from the eastern side of Punta Martin Garcia show the typical duality of dorsal pattern, however, and thus the uniformity of this small series of five is not significant. The large series from Bani, on the other hand, has only a single adult male which shows any pattern; this population is thus almost completely patternless. A fresh series of five topotypes from Fondo Negro has four individuals with patterns, and one without pattern. Thus, although the patternless condition pre- dominates at some localities (Bani), elsewhere (and including the type locality) both types of dorsal pattern occur. The two types of dorsal patterns, as delimited by jMertens, grade into one another. The back may be longitudinally lined with from six to ten tan to yellowish lines of fine dots, or these may be very obscure centrally and more prominent at the sides of the dorsal field. Some individuals have the back finely marbled. The lateral fields are well developed and enclose a series of buffy to cream dots; the lateral fields are often outlined below by a longitudinal yellowish line, and a similar line may border these fields above. The lower sides may be dotted with blue or bluish green. The dotted condition of the back, considered by Mertens as being ahhotti-like, is not at all comparable to the large and brilliant sky-blue spotting on a black ground of that race. There is some similarity between patterned hoekeri and nmhratUis. No nmhratilis however is unpatterned. In patternless lizards, the dorsal ground color was recorded as yellowish brown (Fondo Negro), brown (San Jose de Ocoa), grayish tan (Punta Martin Garcia), black (Barreras), and reddish brown (Bani). The lateral fields may be completely absent or may be indicated by a somewhat grayer longitudinal lateral stripe, without any sort of included or adjacent pale dots. The ventral ground color is blue-gray, purplish blue, blue, light olive, gray, or orange-gray. The throat is likewise variable, but is some shade of dull orange; females have throats which are typically more grayish orange than males. The tails are brown to grayish tan above, and gray below. Scale counts for the series (including intergrades from Hato Nuevo) are: longitudinal ventrals 34-40 (mean 37.2), trans- verse ventrals in 10 (82.4 per cent) or 12 (17.6 per cent) rows, SCHWARTZ AND KLINIKOWSKI : AMEIVA 443 fourth toe scales 73 to 98 (mean 84.6), femoral pores 31-41 (36.5), fifteenth verticil 37-48 (mean 42.7). The much smaller size and faded pattern of those hoekeri which have patterns, as well as the patternless individuals, can easily be distinguished from A. c. chrysolaema. Chrysolaema is likewise characterized by 12 rather than 10 rows of ventrals. In counts of fourth toe scales, femoral pores, and fifteenth verti- cil scales, hoekeri averages less than chrysolaema, the most striking difference being in femoral pores {chrysolaema 43.7, hoekeri 36.5). Patternless hoekeri can be easily differentiated from all umhratilis, since this race is never patternless. Pat- terned hoekeri are much like umhratilis. In both, the dorsal pat- tern is faded and not well demonstrated. One feature is sugges- tive ; patterned hoekeri have the lateral fields prominent and often outlined both above and below, whereas the typical umhra- tilis condition is an obscure lateral field, not set off by longi- tudinal pale lines. Both umhratilis and hoekeri usually have 10 rows of ventrals, although umhratilis has a much higher fre- quency of 12-row individuals. In counts of fourth toe scales and femoral pores, hoekeri averages slightly higher than um- hratilis; the means for fifteenth verticil scales are identical. Because of the similarities of umhratilis and patterned hoekeri, we have considered the possibility that the name hoekeri should be applied to Ameiva from the Yalle de Neiba. To be counted against this conception is the fact that of 61 umhratilis, none is unpatterned, whereas 56.3 per cent of the specimens (hoekeri) from north of the Kio Yaque del Sur are patternless. There is no indication of this patternless condition in specimens from Barahona, nor from elsewhere in the range of umhratilis. We prefer to regard hoekeri as a separate entity, distinct from um- hratilis to the south. A. c. hoekeri is approached geographically by three adjacent races; of these it is known to intergrade only with the race to the northwest in the Valle de San Juan (these intergrades will be discussed later). From umhratilis the range of hoekeri is sep- arated by the lower reaches of the Rio Yaque del Sur and by the extremely mesic conditions of much of the eastern portion of the Valle de Neiba. Although hoekeri is not presently known to intergrade with the race next to the east along the southern coast of the Republiea Dominicana, it may well do so. It is cer- tainly significant that hoekeri occupies the western Llanos de 444 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Azua in the vicinity of Ban! ; just to the east of that city, con- ditions become more mesic, and the region is occupied by another race. Intergradation likely takes place where these two regions come into contact. Specimens examined: Repuhlica Dominicana, Barahona Prov., Fondo Negro, 6 (ASFS X9703-07, MCZ 44757); west side, Punta Martin Garcia, 5 (ASFS V84-88) ; Azua Prov., 3 km E Barreras, 2 (ASFS V3164-65) ; 2 km W Puerto Viejo, 2 (ASFS V3183-84) ; 22 km NW Azua, 3 (ASFS V465-67) ; 1.8 mi. (2.9 km) W, 1.1 mi. (1.8 km) N Azua, 18 (ASFS X8002-18, X8103) ; 1.8 mi. (2.9 km) W, 2.7 mi. (4.3 km) N Azua, 10 (ASFS X8019-28); Peravia Prov., 8.9 mi. (13.9 km) S San Jose de Ocoa, 1300 feet (430 m), 1 (ASFS V714) ; 15.2 mi. (24.3 km) S San Jose de Ocoa, 9 (ASFS V687-95) ; 10 mi. (16 km) NW Bani, 23 (ASFS X7801-21, RT 613-14). Intergrades between A. c. hoekeri and the race to the northwest were examined from : RepuMica Dominicana, Azua Prov., Hato Nuevo, 10 (ASFS X437-46). Ameiva chrysolaema alacris,^ new subspecies Holotype: MCZ 77232, an adult male, from 10 km SE San Juan, San Juan Province, Republica Dominicana, one of a series taken 9 August 1963 by Albert Schwartz and Richard Thomas. Original number V283. Paraiypes: All from the Republica Dominicana, as follows: ASFS V284-97, same data as holotype ; RT 778-79, 10 km S San Juan, San Juan Prov., 9 August 1963, R. Thomas ; MCZ 81005-06, USNM 152561-64, AMNH 92842-43, KU 79865-66, UIMNH 56890-93, 2.5 km W, 4.4 km S San Juan, San Juan Prov., 9 August 1963, D. C. Leber, R. F. Klinikowski ; KU 79867-68, AMNH 92844, 2.5 km W, 5.4 km S San Juan, 9 August 1963, D. C. Leber, R. F. Klinikowski ; ASFS V389-90, 10 km E Valle- juelo, San Juan Prov., 12 August 1963, R. Thomas; USNM 152565, 3 km E Las Matas, San Juan Prov., 9 August 1963, R. Thomas. Associated specimens: Haiti, Dept. du Nord, Cerca-la-Source, 1 (USNM 76780) ; Repuhlica Dominicana, San Rafael Prov., 3.8 mi. (6.1 km) SE Sabana Cruz, 1 (ASFS V329) ; Guayabal, 1 (MCZ 58672); Azua Prov., Tubano (= Padre las Casas), 3 (USNM 66729-31) ; 0.7 mi. (1.1 km) NW Villarpando, 9 (ASFS V419-27). 1 Prom the Latin for "lively." SCHWARTZ AND KLINIKOWSKI : AMEIVA 445 Diagnosis: A subspecies of A. chrysolaema characterized by a combination of moderate size (males to 126 mm, females to 109 mm snout-vent length), usually 10 transverse rows of ventrals, moderate number of fourth toe subdigital scales, very low num- ber of femoral pores, and high number of scales in fifteenth caudal verticil ; dorsal pattern consisting of five to seven bold longitudinal dorsal lines (the lateralmost forming a strong upper border to the prominent black lateral fields with their enclosed bright yellow dots), the longitudinal lines never broken into dots and lines as in c. chrysolaema and always conspicuous and discrete (Fig. 3, left), and a black gular band which rarely in- volves also the chest and undersides of the arms. Fig. 3. Left, Ameiva c. alaeris, holotype, MCZ 77232, 10 km SE San Juau, San Juan Piov., Republica Dominicana. Eight, Ameiva c. proeax, holotype, MCZ 77233, Santo Domingo, 2.2 km SW Rio Ozama, Dist. Nac, Eepublica Dominicana. 446 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY Distribution: From east central Haiti (Cerca-la-Source) south- eastward through the Valle de San Juan (Fig. 11) ; intergrading with A. c. hoekeri at Hato Nuevo, Azua Province, and with A. c. chrysolaema in the vicinity of Mirebalais, Dept. de I'Ouest, Haiti (see discussion below). Description of type: An adult male with the following meas- urements and counts: snout-vent length 116 mm, tail 200 mm; ventrals in 37 longitudinal and 12 transverse rows; fourth toe subdigital scales 42 and 44 (total 86) ; femoral pores 16 and 16 (total 32) ; 46 scales in the fifteenth caudal verticil. Dorsal ground color brown with seven longitudinal pale yellow lines, the lateralmost bordering above the black lateral fields with their isolated yelloAv dots; lateral fields bordered below by a slightly duller longitudinal line which contains a series of bright yellow dots; lower sides dotted with yellow. Throat gray, venter dull, dirtj^ orange. A black gular band, not extending onto the chest or undersides of the arms. Tail brown above, gray below, with an indistinct proximal and lateral area of yellowish dots; top of tail with some darker brown scales. Fore- and hindlimbs with pale scattered small dots, bluish on forelimbs and yellowish on hindlimbs. Variation: See tables. A. c. alacris presents a constant as- semblage of coloration and pattern elements throughout its range. The dorsal ground color is alwaj^s brown, with from five to seven pale or dull yellow longitudinal lines, these lines always forming a conspicuous pattern. The lines are entire and not fragTiiented or modified into series of longitud- inal dots, although in some specimens the more central lines, especially posteriorly, may be broken into dashes. In general, however, the integrity of the lines (even when fragmented) is maintained. The black lateral fields are bold, set off by pale longitudinal lines above and below, and enclose a single series of scattered yellow dots ; the lower line bordering the lateral field may have superimposed upon it a series of bright yel- low dots, thus rendering the black lateral field even more con- spicuous. The lower sides are dotted with yellow. The throat varies from gray to very pale orange, and the venter likewise varies between these two extremes. A. c. alacris intergrades to the southeast with A. c. hoekeri and to the southwest with A. c. chrysolaema. A series of ten specimens from Hato Nuevo, Azua Province (ASFS V437-46), SCHWARTZ AND KLINIKOWSKI : AMEIVA 447 shows the intergradatiou with hoekeri. Of this series, five are un- patterned hoekeri, and five represent the patterned phase of that race. These five patterned lizards have the lateral fields darker than most patterned hoekeri, and there is a distinct tendency to have the dorsal lines more boldly (brighter yellow) displayed anteriorly, although the posterior dorsal pattern is fainter and very like "typical" patterned hoekeri. The distance from Hato Nuevo {alacris X hoekeri intergrades) to Villarpando {alacris) is only ten kilometers, yet the series from the latter locality is typical of alacris in all waj^s and has no patternless individuals. From the vicinity of Mirebalais, Dept. de I'Ouest, Haiti, we have examined specimens from the following localities: 3.4 mi. (5.4 km) NE Barrage de Peligre, 2 (ASFS X2217-18) ; 1.1 mi. (1.8 km) S Mirebalais, 3 (ASFS X2237-39) ; Mirebalais, 1 (MCZ 68510) ; La Tombe, nr. Mirebalais, 8 (MCZ 68517-24) ; Boudou, nr. Mirebalais, 2 (MCZ 69387-88) ; Duvie, nr. Mirebalais, 1 (MCZ 68478). Of these, the last three places named. La Tombe, Boudou, and Duvie, cannot be located ; they have not been mapped. Taken as a whole, this lot of lizards is intermediate between chrysolaema and alacris, although they are closer to alacris than to chrys- olaema. Three lizards (ASFS X2237, ASFS X2217, MCZ 68510) show the disintegration of the dorsal lines into series of yellow spots, a typical chrysolaema feature. Several male specimens are larger than alacris, with snout -vent lengths of 133 to 145 mm (five lizards) ; two females have snout-vent lengths of 109 mm (the upper extreme of alacris females), and another has a snout- vent length of 110. In life, our specimens from Barrage de Peligre and Mirebalais had yellow lines and a greenish wash on the neck — the latter a chrysolaema character. Finally, some individuals have a discrete black gular band as in alacris, whereas others have the band expanded onto the chest and arms as in chrysolaema. We consider this entire lot of specimens inter- gradient between alacris and chrysolaema. Comparisons: A. c. alacris is easily distinguished from the three previously described races on the basis of dorsal pattern ; the discrete, bold, and undotted longitudinal lines of alacris contrast with the patternless or weakly patterned races hoekeri and umhratilis, and with the larger and dorsally dotted and lined chrysolaema. Alacris is a race with ten transverse rows of ventrals as are umhratilis and hoekeri, in contrast to the twelve- rowed chrysolaema. In fourth toe scales, alacris (84.8) averages close to hoekeri (84.6), slightly higher than umhratilis (83.0) 448 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY and lower than chrysolaema (86.7). In femoral pores, alacris has the lowest average (33.8) of any race of A. chrysolaema; of the described forms, it is approached by umhratiUs (35.6) and hoekeri (36.5) and is far below chrysolaema (43.7). In scales in the fifteenth caudal verticil, on the other hand, both alacris and chrysolaema are high (44.0 and 44.4), with hoekeri and unih rat ills (both 42.7) lower. Remarks: A. c. alacris occupies the high and xeric Valle de San Juan and associated upland foothills (i.e., Tubano). It is likely that it is more widespread in east-central Haiti than the present evidence of one specimen from Cerca-la-Source indicates. Presumabl}^ the race follows down the valley of the Riviere de I'Artibonite, and in the vicinity of Mirebalais has genetic con- tact with the more southern chrysolaema. Such genetic con- tinuity may come across the Montagues de Trou d'Eau from the Cul de Sac (although there is no obvious means of penetration of this mountain mass), or chrysolaema may reach Mirebalais via the valley of the Artibonite from the St. Marc area. The only evidence for the latter is the single specimen noted under A. c. chrysolaema, from the "Artibonite Valley"; this individual may have come from some undetermined locality which is inter- mediate between St. Marc and Mirebalais. Further collecting in these areas should easily reveal the precise place of contact between these two races. AmEIVA CHRYSOLAEMA PROCAX^ lieW SUbspCcics Holotype: MCZ 77233, an adult male, from Santo Domingo, 2.2 km SW of the Rio Ozama, Distrito Nacional, Repiibliea Dominicana, one of a series taken 14 June 1963 by Ronald F. Klinikowski, David C. Leber, and Richard Thomas. Original number X7714. Paratypes: All from the Republiea Dominicana, as folloAVS: ASFS X7711-13. X7724-26, MCZ 81007-10, USNM 152566-70, KU 79869-71, UIMNH 56894-97, RT 605-08, same data as holo- type ; ASFS X9254-56, Santo Domingo, old airport, Distrito Nacional, 17 July 1963, D. C. Leber, R. Thomas; AMNH 92845- 49, 5.9 km W Santo Domingo, Distrito Nacional, 20 June 1964. D. C. Leber, R. Thomas. Associated specimens: Rcpuhlica Dominicana, San Cristobal Prov., 8.4 mi. (13.6 km) NE Sabana Grande de Palenque, 2 1 From the Latin for "bold." SCHWAETZ AND KLINIKOWSKI : AMEIVA 449 (ASFS X8167-68) ; 4.2 mi. (6.7 km) NE Sabana Grande de Palemiue, 22 (ASFS X8149-66, RT 643-46). Diagnosis: A subspecies of A. chrysolaenia characterized by a combination of large size (males to 141 mm, females to 116 mm snout-vent length), usually 10 (but often 12) transverse rows of ventrals, moderate number of fourth toe subdigital scales, low number of femoral pores, and high number of scales in the fifteenth caudal verticil ; dorsal pattern a series of six or seven longitudinal yellow lines in a reddish brown field, the lines usu- ally wavy or broken into a series of longitudinal dashes (Fig. 3, right), and a black gular band which may be so expanded as to involve the entire ventral surface including the undersides of the arms. Distribution: The coastal regions of San Cristobal Province and the Distrito Nacional, from the Rio Ozama on the east to the vicinity of Sabana Grande de Palenque on the west (Fig. 11) ; presumed to occur inland, since individuals were seen crossing the road near the city of San Cristobal. Description of type: An adult female with the following measurements and counts : snout-vent length 114 mm, tail 275 mm ; ventrals in 38 longitudinal and 12 transverse rows ; fourth toe subdigital scales 46 and 46 (total 92) ; femoral pores 18 and 20 (total 38) ; 44 scales in the fifteenth caudal verticil. Dorsal ground color rich reddish brown in life, with a series of seven dull longitudinal lines, the median line rather obscure, the lateral lines broken into a series of wavy dashes, the lateralmost lines bordering above the black lateral fields, which have a series of tiny yellow dots inclosed within them. Sides of head gray with a creamy subocular patch and some pale irregular blue blotches. Throat fleshy gray. Ventral color dark blue-gray. Hindlimbs heavily spotted with yellow, forelimbs faintly spotted with blue. A dark gray gular band not involving the chest and underside of the arms. Tail reddish brown above, dark gray below. Variation: See tables. The series of A. c. procax is remark- ably uniform in both coloration and pattern. The dorsal ground color is always some shade of reddish brown, and there may be a yellowish green wash over the shoulders. The longitudinal lines are conspicuous, although the median line may be reduced or faint. Only in young and subadult individuals are the lines entire, and even in these specimens there is a strong tendency toward wavy fragmentation. The lateral fields are black and the yellow dots included therein are regularly very tiny; in 450 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY females these included dots are less well differentiated than in the males. The gular band is present, and in large males may be very extensive, including most of the belly. The venter is blue in juveniles and females, and gray to solid black in males. Comparisons: The reddish brown dorsal coloration and the wavy dorsal lines separate procax from all other races. Some specimens of chrysolacma were noted in life as reddish brown, but this is not the usual dorsal coloration of the nominate race. In- dividuals of chrysolaema with this coloration are readily differen- tiated from procax on the basis of the very different dorsal pat- terns of the two subspecies. In size, procax is closest to chrysolaema, although procax is dis- tinctly the smaller of the two. Chrysolacma is typically a subspe- cies with 12 rows of ventrals, whereas procax, although it has in- dividuals with 12 rows, has a modal condition of 10 rows. Procax averages fewer (84.8) fourth-toe scales than chrysolaema (86.7), the same as alacris, and more than hoekeri (84.6) and umhratilis (83.0). In number of femoral pores, procax (36.1) averages far lower than chrysolacma (43.7), and slightly lower than hoekeri (36.5), but slightly higher than umhratilis (35.6) and alacris (33.8) ; procax is one of the races with a low^ number of femoral pores. In fifteenth verticil scales, procax averages less (42.8) than chrysolaema (44.4) and alacris (44.0), and is about equal to hoekeri and umhratilis (42.7). Remarks: A. c. procax is presently not known to intergrade either with hoekeri to the west or the race next to the east along the southern Hispaniolan coast. The easternmost hoekeri locality (where the population incidentally is almost completely made up of non-patterned individuals) and the westernmost locality of procax are separated by onh' 35 kilometers. As noted in the dis- cussion of hoekeri, Bani lies about on the dividing line between the xeric Llanos de Azua to the west and more mesic conditions on the east. Procax inhabits these eastern more mesic regions, whereas hoekeri is restricted to the xeric Llanos de Azua. Ameiva CHRYSOLAEMA PARVORis,^ new subspecics Holotype: MCZ 77234, an adult male, from 0.9 mi. (1.4 km) E Boca Chica, Distrito Naeional, Republica Dominicana, one of a series taken 23 x\ugust 1963 by Ronald F. Klinikowski, David C. Leber, and Richard Thomas. Original number V649. 1 From the Latin parvum (small) and os, oris (mouth), a translation of Boca Chica. the type locality. SCHWARTZ AND KLINIKOWSKI : AMEIVA 451 Paratypes: All from the Republica Dominicana, as follows: MCZ 81011-14, USNM 152571-74, A:MNH 92850-55, KU 79872-74, UIMXH 55638-39, RT 789-90, same data as liolotype; ASFS V669-79, Boca Cliica, eastern edge, Distrito Nacional, 23 August 1963, R. F. Klinikowski, D. C. Leber, R. Thomas. Associated specimens: Repuhlica Dominicana, San Pedro de Macoris Pro v., 0.5 mi. S San Pedro de Macoris, at lighthouse, 11 (ASFS X8181-91) ; La RomanaProv., Isla Catalina, western end, 4 (ASFS V554-57). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of large size (males to 137 mm, females to 113 mm snout-vent length), usually 12 transverse rows of ventrals, mod- erate number of fourth-toe subdigital scales and femoral pores, and high number of scales in the fifteenth tail verticil; dorsal pattern of tan to blackish brown ground color with dull yellow spots which may be either discrete or confluent, giving a reticu- late appearance (Fig. 4, left), lateral fields present and black, or broken to give a tigroid effect, and a black gular band which may be expanded to involve the chest and the undersides of the arms. Distribution: Coastal southeastern Republica Dominicana, from Boca Chica on the east to San Pedro de Macoris on the west, and including Isla Catalina ; range as here described apparently discontinuous, and A. chrysolaema unknown from the mainland opposite Isla Catalina (Fig. 11). Description of type: x adult male with the following meas- urements and counts : snout-vent length 129 mm, tail 282 mm ; ventrals in 38 longitudinal and 10 transverse rows; fourth-toe subdigital scales 41 and 42 (total 83) ; femoral pores 19 and 19 (total 38) ; 44 scales in the fifteenth tail verticil. Dorsal ground color dull blackish brown, with the entire back from the neck to the sacrum covered with dull yellow spots, not aligned into linear series ; lateral fields black, not bordered above or below, and in- vaded by the brown dorsolateral coloration, the entire sides spot- ted with yellow dorsally and pale blue ventrally. Throat gray, ventral ground color gray. A black gular band which extends slightly onto the chest and also the underside of the arms. Fore- limbs faintly spotted with bluish, hindlimbs spotted with dull yellow. Tail grayish brown dorsally, gray ventrally. Variation: See tables. The dorsal ground color of A. c. par- voris varies from blackish brown (type locality) to tan and brown (San Pedro de Macoris). The spotted condition of the black is typical of most specimens, including small juveniles. 452 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Fig. 4. Left, Ameiva c. parvoris, holotype, MCZ 77234, 0.9 mi. E Boca Chica, Dist. Xac, Eepublica Domiuicana. Eight, Ameiva c. jacta, holotype, MCZ 75267, Juanillo, La Eouiana Prov., Eepublica Dominicana. but others show a more lineate pattern somewhat like that described for procax — the dorsal lines broken into dashes giving a wavy appearance. The lateral fields maj^ be obscured as in the type, or may be slightly more prominent, with much encroachment of brown to give an irregular and indefinite edge ; at the same time the black lateral field pigment may extend up onto the sides of the back, thereby giving a distinctly tigroid appearance to the sides and dorsolateral regions. In some indi- viduals the dorsal spots are confluent, thereby increasing the tigroid effect by transverse dorsal pale markings. The throat is SCHWARTZ AND KLINIKOWSKI : AMEIVA 453 grayish to dull orange and the ventral ground color varies from grayish blue to dull, deep orange. The small series from Isla Catalina resembles the mainland specimens in dorsal pattern and in extent of the gular band, which in parvoris may involve the chest and undersides of the arms. The dorsal dots are conspicuously confluent, the lateral fields are obsolete, and the ventral coloration is pale bluish with an orange tint. The most obvious difference in coloration is that the dorsal surfaces of the hindlimbs are rusty — a feature found in no mainland parvoris. The Catalina series is composed of one adult male with a snout-vent length of 126 mm and thus a large lizard, and three females (one of which is a small juvenile), the largest of which has a snout-vent length of 112 mm, again a large lizard. The transverse ventrals are 10 in three specimens and 12 in one. In all other features of scalation the Catalina lot falls within the known range of mainland parvoris. Additional speci- mens from Isla Catalina ma}- well reveal that it is inhabited by still another distinctive race ; the strikingly rusty hindlimbs are indicative of at least one major color difference between Isla Catalina specimens and mainland parvoris. Comparisons: No other race of A. chrysolaema has the back with irregularly arranged and at times confluent spots, and tigroid sides. This feature alone will distinguish parvoris from the previous races. In size, parvoris is much smaller than chryso- laema and slightly smaller than procax, but larger than the re- maining subspecies. In fourth toe scales, parvoris averages less (83.2) than other races except umhratilis, which is comparable (83.0). Pari^om averages less (38.2) than chrysolaema (43.8) in femoral pores, but exceeds the remaining races, all of which have low femoral pore counts. In fifteenth verticil scales, parvoris has the same mean as hoekeri and umhratilis (42.7), less than chry- solaema and alacris (44.4 and 44.0), and almost the same as procax (42.8). Remarks: A. c. parvoris is not known to intergrade with procax on the west nor with the following subspecies to the east. The easternmost locality for procax is separated by 35 kilometers from the westernmost parvoris record. It is possible that the Rio Ozama may divide these two races of A. chrysolaema. The area occupied by parvoris does not differ in any obvious way from that inhabited by procax. 454 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Ameiva chrysolaema jacta^ new subspecies Holotype: MCZ 75267, an adult male, from Juanillo, La Ro- mana Province, Republica Dominicana, taken 29 March 1963, by Clayton E. Ray and Robert R. Allen. Paratypes: MCZ 75268-69, same data as holotype. Diagnosis: A subspecies of A. chrysolaema characterized by a combination of large size (males to 134 mm snout-vent length; females unknown), usually 12 transverse rows of ventrals, low number of fourth toe subdigital scales, moderate number of femo- ral pores, and very low number of scales in the fifteenth caudal verticil; a dorsal pattern of dark brown to black tigroid mark- ings on a grayish tan background (in preservation), the pattern extending in a diluted fashion onto the neck, a bold, checker- board-patterned tail (Fig. 4, right), and a black gular band which extends onto the chest and underside of the arms. Dist)ihidion: Known only from the type locality in extreme eastern Republica Dominicana (Fig. 11). Description of type: An adult male with the following meas- urements and counts : snout-vent length 134 mm, tail 304 mm ; ventrals in 38 longitudinal and 12 transverse rows; fourth-toe subdigital scales 41 and 41 (total 82) ; femoral pores 21 and 22 (total 43) ; 35 scales in the fifteenth caudal verticil. Dorsum (preserved) tannish gray with a dark brown, almost black pat- tern consisting of bold vertical tigroid markings on the sides and about five wide, dark, longitudinal lines on the back, the latter much confused and joined by the lateral vertical markings, giv- ing a rather complete and complex dark brown reticulum, which, although present on the neck, is much paler gray. Lateral fields completely absent, no dotting on sides or back whatsoever. Upper surface of forelimbs with obscure grayish lines and blotches, hindlimbs with a dark brown reticulum enclosing large spots which are pale centrally and darker peripherally. Tail with a bold checkerboard pattern of grayish tan, white, and dark brown, this pattern becoming obsolescent and absent on the distal half of the tail. Belly and throat presently dull grayish orange, with some lateral ventral scales with very dark gray blotches, thereby giving the belly somewhat of a faint checkerboard appearance laterally. Black gular band obsolete but indicated, and anterior ten rows of ventrals clouded with dark gray. 1 From the L.itin for "thrown," an nllusion to the far flung distribution of this subspecies. SCHWARTZ AND KLINIKOWSKI : AMEIVA 455 Variation: See tables. In coloration and pattern the two paratypes are much like the type and require little comment. The major difference is that the paratypes have a bold, black, gular band which involves the chest and the underside of the arms. Neither lizard has any indication of lateral fields, and the sides and back have the tigroid vertical bars and brown to black reticulum, as well as the checkerboard tail, just as de- scribed for the type. Comparisons: A. c. jacta does not need detailed comparison of pattern with any other described subspecies ; the boldly and viv- idly marked back with its light ground color will distinguish jacta from the remaining races. A. c. parvoris is closest to jacta in pattern, but the differences are so striking that the similarity between these two subspecies is not very great. In size, jacta is smaller than chrysolaema, procax and parvoris, and larger than the remaining forms. All other races have a higher numl^er of fourth-toe scales and fifteenth verticil scales. Jacta has a high mean of femoral pores, having less than chryso- laema, and more than the balance of the subspecies. Remarks: No intergradation is known between jacta and par- voris to the southwest; the easternmost mainland locality for parvoris is separated by 145 kilometers from that of jacta. We have attempted to secure specimens of A. chrysolaema between San Pedro de Macoris and Juanillo at several localities along the coast (La Romana, Boca de Chavon, Boca de Yuma) as well as inland in this eastern region, without success. Typical xeric chry- solaema habitats here are occupied by A. taeniura. Considering the likeness of jacta to the race from Isla Saona, described below, it is probable that this boldly marked type of lizard was at one time (and still is?) abundant locally in extreme eastern Hispan- iola. Presently, the hiatus between jacta and parvoris and the apparent absence of the species in this eastern region suggests strongly that the populations are relict wdth a disjunct distribu- tion. AmEIVA CHRYSOLAEMA RICHARDTHOMASI ^ UCW SUbspecicS Holotypc: :\ICZ 77235, an adult male, from the environs of Mano Juan, Isla Saona, Repiiblica Dominicana, taken 19 July 1964 by Richard Thomas. Original number V3018. 1 Named for the collector. 456 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY Paratypes: ASFS V3019-30, USNM 152575-76, AMNH 92856- 58, KU 79875-77, UIMNH 56898-99, RT 935, same data as holo- type. Diagnosis: A subspecies of A. chrysolaema characterized by a combination of large size (males to 137 mm, females to 124 mm snout-vent length), usually 10 (but often 12) transverse rows of ventrals, high number of fourth-toe subdigital scales and femoral pores, and moderate number of scales in the fifteenth caudal ver- ticil; dorsal pattern of two phases: (1) back gray-green with Fig. 5. Left, Amelia c. richardthomasi, holotype, MCZ 77235, environs of Mano Juan, Isla Saona, Eepublica Dominicana. Eight, Ameiva c. ridtard- thomasi, ASFS Y3019, environs of Mano Juan, Isla Saona, Eepublica Dominicana. SCHWARTZ AND KLINIKOWSKI : AMEIVA 457 only an indistinct mottling of gray-brown in the area of the lat- eral fields or (2) back gray-green with heavy black mottling, no lateral fields, and tigroid vertical bars on the sides, the dorsal mottling not extending onto the shoulders and neck (Pig. 5), and without a black gular band. Di.strihufion: Known only from the type locality, but pre- sumed to occur throughout Isla Saona (Fig. 11). Description of type: An adult male with the following meas- urements and counts : snout-vent length 137 mm, tail 146 mm, twice broken and regenerated ; ventrals in 37 longitudinal and 12 transverse rows ; fourth-toe subdigital scales 45 on left side ; femoral pores 22 and 20 (total 42) ; 38 scales in the fifteenth tail verticil. Dorsal ground color gray-green (Maerz and Paul: pi. 22F1), becoming finely mottled laterally with a series of \Qvy faint grayish brown vertical bars in the region of the lateral fields ; lower sides putty colored. Throat and venter orange with no black gular band. Fore- and hindlimbs unicolor with back, and patternless. Tail grayish tan without obvious checkerboard pattern above, putty colored below. Variation: See tables. Sixteen specimens of richardthomasi (including the type) are patternless dorsally and have a finely filigreed, grayish brown area in the region of the lateral fields. The tails are gray to tan without a prominent checkerboard pat- tern as in the patterned lizards. The venters are orange to drab gray, generally slightly more orange in the pectoral region, and grayer posteriorly. Throat and underside of forelimbs are mottled to nearly unicolor orange, sometimes in discrete flecks. The younger specimens have light gray throats. The underside of the hindlimbs and tail is gray to putty colored with some orange on the anterior surface of the femur, and in some speci- mens on the entire underside of the hindlimb. Two small juve- niles (snout-vent length 47 and 51 mm) are in this patternless phase. Eight specimens (adults and subadults of both sexes) have patterned backs. In this phase, the dorsal ground color is gray- green with a heavy black mottling and strikingly' tigroid barred sides. The black dorsal x^attern quickly fades at the shoulders and is absent or very suppressed on the neck. The dorsal blotch- ing is much as that described for jacta, i.e., a vermiculate or at times longitudinally arranged configuration of black on a lighter ground. In no specimens are the lateral fields apparent and there is no spotting or dotting on the sides. The bellies of these 458 BULLETIN: MUSEUM OF COMPARATR^ ZOOLOGY patterned lizards were the same as those of patteruless speci- mens; females of the patterned lizards have the extent and in- tensity of the orange not so great as do the males. The checker- board tail is a common feature. As in A. c. hoekeri, we have no doubt that the two phases in richardthomasi represent two basic patterns, and in no way should be interpreted as an adult phase versus a juvenile and subadult phase. Although the only two juveniles at hand are patternless, there is an intermingling of sizes of both sexes insofar as the two phases are concerned. Just as in hoekeri, which is rep- resented by a much longer series, there are no intermediates be- tween the two conditions; each lizard is distinctly in one phase or the other . Comparisons: The patterned phase of A. c. richardthomasi re- quires comparison only with A. c. jacta to which race the former is obviously closely allied. The two can easily be differentiated in that jacta has a black gular band and the dorsal pattern con- tinues anteriorly onto the neck, whereas richardthomasi lacks a gular band and has the pattern faded anteriorly. Jacta also is not known to have a patternless phase The patternless phase of richardthomasi requires comparison with patternless hoekeri. The two are much alike, but richard- thomasi differs in having the filigreed or mottled lateral field area whereas hoekeri has an obsolete lateral field and no mottling in this region. Also the dorsal hues of hoekeri populations are usually not greenish. In size, richardthomasi is smaller than chrysolaema and procax, equal to parvoris, and larger than the remaining races. In having a mean of 87.6 fourth-toe scales, richardthomasi averages higher than all previously named races. In femoral pore counts, it is higher than all races except chrysolaema. Considering fifteenth caudal verticil scales, richardthomasi is exceeded by all forms ex- cept jacta. Remarks: The close alliance of richardthomasi with jacta is obvious. If we assume that extreme eastern Hispaniola is (was) inhabited by a population with heavy dorsal mottling and marbling, this population must also have given rise to the Saonan subspecies. We have no evidence that jacta or a related form still occurs on the adjacent Hispaniolan mainland. Certainh' richardthomasi is an insular derivative of an extreme eastern heavilv marked form which was likelv similar to jacta. SCHWARTZ AND KLINIKOWSKI : AMEIVA 459 Ameiva chrysolaema leberi^ new subspecies Holohjpe: MCZ 77236, an adult male, from 5 km E Peder- nales, Pedernales Province, Republica Dominicana, one of a series taken 25 June 1964 by David C. Leber and Richard Thomas. Original number V2509. Paratypes: All from the Republica Dominicana, Pedernales Prov., as follows: ASFS V2510-19, USNM 152577-82, RT 932, KU 79878-81, same data as holotype; MCZ 81015-18, UIMNH 56900-05, Pedernales, 3 July 1964, D. C. Leber, R. Thomas; AMNH 92859-61, 1 km E Pedernales, 25 July 1963, R. F. Klini- kowski ; AMNH 92862-63, 12 km E Pedernales, 25 June 1964, R. Thomas. Associated specimens: Haiti, Dept. d l' Quest, Tean, nr. Sal- trou (not mapped), 4 (MCZ 69389-92) ; Saltrou, 7 (AMNH 50000- 04, 50007-08). BepuMica Dominicana, Pedernales Prov., Oviedo, 3 (MCZ 58674-76). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of small size (males to 111 mm, females to 104 mm snout-vent length), 10 transverse rows of ventrals, moderate num- ber of fourth-toe subdigital scales and femoral pores, and low number of scales in the fifteenth caudal verticil ; a completely pat- ternless rusty brown dorsum, no lateral fields (Pig. 6, left), a deep orange-red belly, and a black gular band which may involve the chest and underside of the arms. Distribution : To the south of the Massif de la Selle and Sierra de Baoruco, from the vicinity of Saltrou in Haiti, east onto the Peninsula de Barahona, to 12 kilometers southeast of Pedernales (Fig. 11). The record from Oviedo is discussed below. Description of type: An adult male with the following meas- urements and counts : snout-vent length 103 mm, tail 233 mm ; ventrals in 34 longitudinal and 10 transverse rows; fourth-toe subdigital scales 40 and 42 (total 82) ; femoral pores 21 and 20 (total 41) ; 40 scales in fifteenth caudal verticil. Dorsum uniform rusty brown anteriorly, becoming gray-brown posteriorly; sides of head reddish brown. Lateral fields absent. Throat orange, ven- tral ground color brick red, lower sides (and lateralmost two rows of ventral scales) blue. A black gular Imnd, extending onto the first four or five rows of ventrals and onto the undersides of the arms. Tail gray above, off-white below. Dorsum and top of tail completely unpatterned. 1 Xamed for one of the collectors. 460 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Fig. 6. Left, Ameiva c. leheri, holotype, MCZ 7720(5, '> km E Pederualcs, Pedeniales Prov., Kepubliea Dominicana. Bight, Ameiva c. ficta, holotype, MCZ 77237, 13.1 mi. SW Euriquillo, Pedernales Prov., Eepubliea Dominicana. Variation: See tables. The dorsal ground color varies from rusty brown to reddish brown, and the ventral ground color from gray with small amounts of red to orange-red, fading posteriorly, to brick red. The gular band is present in all specimens, and in only one male does it not extend onto the chest. The lower sides and lateral two rows of ventrals on each side are blue, or at least have blue flecking, but one small male has the lateral ventrals orange-red like the balance of the venter. The lores and cheeks are unspotted pinkish gray. The tails are gray, unmarked above, SCHWARTZ AND KLINIKOWSKI : AMEIVA 461 and the undersides of the tails are a grayish off-white. There is no obvious sexual dichroniatism. Comparisons: By virtue of its patternless dorsum, leheri can easily be differentiated from all races except patternless hoekeri and patternless richardthomasi. The different dorsal hues of leheri and ricJiardfhomasi (rusty brown versus greenish gray) and the presence of a gular band in the former and its absence in the latter, as well as the larger adult size of richardthomasi, all make this distinction easy. From patternless hoekeri, leheri dif- fers in smaller adult size, and in lacking any expression of the lateral fields, whereas boekeri retains the fields as obsolete grayish longitudinal smudges. The vivid venters of leheri are not found in hoekeri, and the brighter dorsal ground color of leheri con- trasts strongly with the more drab tones of hoekeri. Leheri is the smallest race of A. chrysolaema. In number of fourth-toe scales, leheri averages lower (85.5) than richard- thomasi (87.8) and chrysolaema (86.7), and is higher than the remaining subspecies. In femoral pores, leheri again averages less (41.3) than chrysolaema (43.7) and richardthomasi (42.6), the same as jacta, and more than in the other races. In fifteenth verticil scales, leheri is exceeded by all subspecies except jacta. Remarks: A. c. leheri is not known to intergrade with either chrysolaema to the north (from whose range it is completely separated by the Massif de la Selle) or with the form to the east on the Peninsula de Barahona. The three specimens from Oviedo noted in "Associated Specimens" above, will be discussed in detail in the treatment of the following subspecies. AmeIVA CHRYSOLAEMA FICTA^ UCW SUbspCcicS Holotype: MCZ 77237, an adult male, from 13.1 mi. (20.8 km) SW Enriquillo, Pedernales Province, Republica Dominicana, one of a series taken 22 July 1963 by Albert Schwartz and Richard Thomas. Original number X9401. Paratypes: All from the Repiiblica Dominicana, Pedernales Province, as follows: ASFS X9402-09, same data as holotype; ASFS X9950, same locality as holotype, 30 July 1963, R. Thomas; ASFS V197-98, same locality as holotype, 4 August 1963, D. C. Leber, R. Thomas. Associated specimens: Repuhlica Dominicana, Pedernales Prov., 30 km from Oviedo, road to Pedernales, 1 (MCZ 58673) ; 1 Prom the I-atin for "invented, devispfl," in allusion to resemblances to abbntti. 462 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Oviedo, 15 (MCZ 58677-80, 58682-90, 58692-93) ; 5 mi. (8 km) NE Oviedo, 11 (ASFS X9957-58, V273-80, RT 752) ; Barahona Prov., 3 km SW Enriquillo, 1 (ASFS V290) ; Enriquillo, 2 (MCZ 58777-78). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of moderate size (males to 121 mm, females to 113 mm snout-vent length), 10 transverse rows of ventrals, low number of fourth-toe subdigital scales, moderate number of femoral pores and scales in the fifteenth tail verticil; dorsum tan to brown, spotted with pale blue, lateral fields obsolescent and often replaced by vertical blackish bars or vermiculations (Fig. 6, right), belly rust colored, a prominently checkerboarded tail, and a black gular band often extending onto the chest and underside of the arms. Distribution: The Peninsula de Barahona from (apparently), 30 km NW Oviedo in the west, east to the east coast in the vicin- ity of Oviedo, and thence north to Enriquillo (Fig. 11) ; see however Remarks below. Description of type: An adult male with the following meas- urements and counts : snout-vent length 112 mm, tail 287 mm ; ventrals in 36 longitudinal and 10 transverse rows; fourth toe subdigital scales 38 and 39 (total 77) ; femoral pores 21 and 20 (total 41) ; 38 scales in the fifteenth caudal verticil. Dorsal ground color brown, with six longitudinal series of more or less discrete pale blue spots, the lateralmost series bordering above the remnants of the lateral fields, below which is another longi- tudinal series of pale blue spots; lower sides with alternating vertical black and pale blue bars ; f orelimbs vaguely spotted with small dots, hindlimbs boldly marked Avith large rusty spots dorsally. Throat grayish orange, belly rust colored; an exten- sive black gular band which includes the first eight rows of ventrals and extends onto the undersides of the arms. Tail brown above, with blue spots on the first nine verticils dorsally, and additionally somewhat checkerboarded; tail ivory below. Variation: See tables. The tan to brown dorsum with pale blue discrete spots characterizes A. c. ficta. The spots, on occasion, may be greenish anteriorly or tan posteriorly, but in general they are pale blue. The lateral fields are obsolete or almost completely obliterated by vertical black bars alter- nating wdth blue bars on the sides. The throat and ventral ground color are grayish orange and rust, with bellies of females slightly paler than those of males. In some individuals the dorsal spots are distinctly lineate in appearance, and in a few SCHWARTZ AND KLINIKOWSKI : AMEIVA 463 the back has a more or less complete finely filigreed appear- ance, although this is not the norm. The spots themselves vary in size, distribution, and density ; they may be much smaller than in the type and much more closely appressed to one an- other, or the paramedian rows (if rows are discernible) may be fused to form a pair of paramedian pale blue lines. The checkerboard tail with blue spotting on its basal portion is a common feature. Comparisons: Only one other race thus far discussed, A. c. parvoris, has a spotted dorsum, although A. c. chrysolaema has a pattern of spots and lines. In neither of these two races are the dorsal spots pale blue, but are rather some shade of yellow. In actuality, the pattern of chrysolaema, although dotted, bears little resemblance to that of ficta; the pattern of parvoris is similar but the coloration and general aspect of the lizards of these two races are quite distinctive. Parvoris lacks a conspicu- ously checkerboarded tail. Compared to the described races, jicta is exceeded in size by all other forms except leheri, which is still smaller. In number of fourth-toe scales, ficta is exceeded by all races except jacta. Chrysolaema, richardthomasi, jacta, and leheri exceed ficta in mean number of femoral pores, and in this character ficta exceeds the balance of the races. Ficta averages higher in fifteenth verticil scales than richarclthomasi, leheri, and jacta, and lower than the other races. Remarks: The distribution of A. c. ficta encompasses the east- ern shore of the Peninsula de Barahona from Enriquillo south to Oviedo, and thence inland toward Pedernales for a distance of 30 kilometers. Ficta is not known to intergrade with either imibratilis to the north or leheri to the west (but see below). The northernmost station for ficta is 40 kilometers from the closest record of umhratilis ; we presume that these two races do not come in contact because of the, at best, narrow and intermittent nature of suitable habitats for chrysolaema along the east coast of the Peninsula de Barahona. There are three specimens (MCZ 58674-76) from Oviedo which are clearly leberi and in no way resemble ficta. Assuming that these specimens did indeed come from Oviedo, they present a problem. They are the only specimens from the entire eastern coast of the Peninsula de Barahona which are patternless ; in our considerable collecting experience in the Oviedo region, we never encountered nor collected any leheri-\ike individuals. There are 464 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY several possibilities ; all of which depend upon the assumed cor- rectness of the locality data for these three specimens: 1) they represent a Ze&en-phase of ficta; 2) leheri and ficta are not both subspecies of chrysolaema; 3) leheri and ficta are both chrysolaema derivatives but one has been so long separated from the parent stock that the two forms act as species, with a rather broad (30 km) region of sympatry. In defense of our arrangement of these two forms as subspecies of A. chrysolaema, the following comments are pertinent: 1) There is no incontrovertible evidence that these three lizards are a patternless phase of ficta. All of our own Oviedo material is pat- terned, and there is no indication that there exists a patternless phase oi ficta (although of course there is this possibility). 2) Since neither Ichcri nor ficta intergrades with any other subspecies for reasons of geography, and since both have apparently been long isolated from chrysolaema and umhraiilis to the north, one or both might be logically regarded as a distinct species (if so, then ahbotti and ficta would compose one species, or leheri could be so regarded). We feel that leheri, despite its complete isolation from chrysolaema, is so like patternless boekeri and richard- thomasi that to regard it as a distinct species would be mislead- ing and obscure its clear relationships to the balance of A. chrysolaema. A somewhat stronger case may be made for sep- arating ahhotti and ficta at the species level; here again, how- ever, the resemblance of both these spotted forms to parvoris for example (as well as the overall similarities of ahhotti- ficta to the more northern subspecies) tends in our opinion to negate removing these two forms from the species chrysolaema. 3) The most appealing interpretation is that one (leheri) of the two in- volved forms has been long separated from its parent stock (A. c. chrysolaema), and that once contact between it and another subspecies (ficta) has been re-established, the two forms do not intergrade but act as separate species. The present lack of contact between leheri and chrysolaema and between ficta and umhratilis suggests that both forms may well have had long inde- pendent histories. It is even not improbable that ficta has been derived from aMotti, rather than the reverse, and thus ahhotti may have been insularly isolated from leheri. Such a combination of situations might argue for species status for both leheri and ahhotti-ficta and we have considered this possibility. On the other hand, such a decision obscures the obvious relationships of these two forms to A. chrysolaema (in contrast, for instance, to A. taeniura or A. lineolata) . SCHWARTZ AND KLINIKOWSKI : AMEIVA 465 Finally, and probably the most important point is that the region between Oviedo and Pedernales still remains little known herpetologically ; there is alwaj^s the possibility that the pre- sumed leheri from Oviedo are in actuality from farther west and thus from within the known range 0/ leheri. We have tried to adhere in this ease to a via media, and rather than make assump- tions from inadequate data, we consider both leheri and ficta sub- species of A. chrysolaema, although admitting that the situa- tion is not completely clear. Exclusive of these three question- able lizards, the ranges of jicta and leheri approach, very closely; the distance between the nearest localities for the two races is only 15 kilometers. AmEIVA CHRYSOLAEMA ABBOTTI Noble, 1923 Ameiva ahhotti Noble, 1923, Amer. Mus. Novitates, 64:1 (type locality — Isla Beata, Eepublica Dominicana). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of small size (males to 117 mm, females to 108 mm snout-vent length), usually 12 transverse rows of ventrals, high number of fourth-toe subdigital scales and femoral pores, and moderate number of scales in the fifteenth verticil; dorsum black with a pattern of isolated spots which are orange or yellowish, becoming blue anteriorly, lateral field absent, the sides spotted with sky-blue spots (Fig. 7, left), venter deep brick red to orange posteriorly, and a black gular band which expands to cover the chest and underside of the arms. Distrihution: Known only from Isla Beata, off the tip of Cabo Beata (Fig. 11). Disciissio7i: A. c. ahhotti is the most brilliantly colored and striking of the races of A. chrysolaema. The combination of black dorsal coloration, orange to yellowish spots middorsally, becom- ing blue anteriorly, and the vivid blue lateral spots provides a particularly colorful lizard. The forelimbs are black to brown (distally) with blue spots, and the hindlimbs black with prox- imally blue and distally orange spots. The venter is brick red, grading to orange or pinkish posteriorly, and the lateralmost ventrals are invaded by blue and white spotting. The heads are tan to orange with white or bluish spots on the sides. There is a bold black pectoral band which extends onto the chest and even onto the venter and the undersides of the arms. The underside of the hindlimbs is orange on the thighs and orange 466 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Fig. 7. Left, Ameiva c. abbotti, ASFS V2743, Isla Beata, Eepublica Dominicana. Eight, Ameiva c. secessa, holotype, MCZ 77238, Etroits, He de la Gonave, Haiti. to light tan on the crura. The upperside of the tail is checker- boarded black and white or cream; the underside of the tail is gray to greenish on its proximal half to two-thirds, and uniform light tan to pinkish distally. The dorsal spots are invariably discrete and there is neither a tendency for them to become lineate or to be fused into longitudinal lines. The females are colored and patterned like the males; a juvenile lacks the bright orange ventral color and the black pectoral area. A. c. ahhoiti is so very distinctive in its dorsal coloration and pattern that it is hardly necessary to compare it with any other SCHWARTZ AND KLINIKOWSKI : AMEIVA 467 subspecies. Closest, at least in pattern, are parvoris and ficta; the former, although a dorsally spotted race, is not so gaudy and lacks the discrete spotting so characteristic of abhotti. The ad- jacent race ficta, on the mainland, resembles ahhotti in basic pattern, but differs in having the back brown rather than black, in having the dorsal spots pale blue rather than orange to yellowish, in having the spots at times arranged into lines, and in having the spotting on the back quite variable in density. In contrast, ahhotti is surprisingly constant in density of dorsal spotting. Ficta is primarily a race with 10 rows of ventrals. whereas ahhotti usually has 12. Variation: See tables. A. c. ahhotti has the highest mean (89.6) of fourth-toe scales of any race described to this point, and is ap- proached only by richardthomasi (87.8) ; of all races of A. chry- solaema, ahhotti has the highest average of femoral pores (43.8), although it is closely approached by chrysolaema (43.7). AVith a mean of 40.4 fifteenth verticil scales, ahhotti exceeds ficta, richard- thomasi, leheri and jacta, and has less verticil scales than the other races. The relationships of abhotti are obviously with ficta on the mainland. Whether the latter occurs on the southern tip of the Peninsula de Barahona is unknown, but it certainly is a more likely candidate there than the drab and patternless leheri, if we assume that abhotti was derived directly from the adjacent mainland. Specimens examined: Repuhlica Dominicana, Isla Beata, 39 (ASFS V2743-69, MCZ 28571-73, 37578-79, 37581-82, 17676-77, 57049, RT 934, UMMZ 83098). Ameiva chrysolaema secessa^ new subspecies Holotype: MCZ 77238, an adult male, from Etroits, He de la Gonave, Haiti, taken 17 July 1962 by Elie Cyphale. Original number X2447. Paratypes: All from He de la Gonave, as follows: ASFS X2440-46, X2448-59, UIMNH 56906-09, USNM 152583-87, AMNH 92864-69, KU 79882-86, same data as holotype ; USNM 80377-78, Pointe Quest, 21 March 1930, L. H. Parish and W. Perrygo; USNM 77062-69, MCZ 25539-48, Pointe a Raquette, August 1927, W. J. Eyerdam; MCZ 80251-78, Pointe a Raquette, sum- mer 1964, G. Whiteman; MCZ 80231-36, Nan Palmiste, 4 km 1 From the Latin for "distant, removed." 468 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY from Pointe a Kaquette, summer 1964, G. Whiteman; MCZ 80237-50, Ti Roche, 0.5 km from Pointe a Raquette, summer 1964, G. Whiteman; USNM 80359-68, 80370-76, UMMZ, 92196, Anse a Galets, 23 March 1930, L. H. Parish and W. Perrygo; MCZ 37568-77, Anse a Galets, 9 April 1934, T. Barbour; USNM 76803, Nan Cafe, March 1929, A. J. Poole and W. Perrygo. Associated specimens: He de la Gondve (no other locality), 3 (CM 8133, MCZ 12870-71). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of large size (males to 134 mm, females to 111 mm snout-vent length), usually 12 transverse rovs^s of ventrals, high number of fourth-toe subdigital scales, and moderate number of femoral pores and scales in the fifteenth caudal verticil ; dor- sum reddish brown with six or seven dull buffy longitudinal lines, grayish brown (rather than black) lateral fields with isolated buffy dots enclosed therein (Fig. 7, right), and with the black gular band usually absent, or at least very restricted, seldom involving the anterior ventrals but at times extending onto the underside of the arms. Distribution: He de la Gonave, Haiti (Fig. 11). Description of type: An adult male with the following meas- urements and counts: snout-vent length, 117 mm, tail 134 mm, broken; ventrals in 39 longitudinal and 10 transverse rows; fourth-toe subdigital scales 45 and 45 (total 90) ; femoral pores 21 and 21 (total 42) ; 42 scales in the fifteenth caudal verticil. Dorsal ground color reddish brown with seven dull buffy longi- tudinal lines, the median line somewhat broken and indistinct; head dull tan, neck greenish ; sides of head gray with whitish blotches. Lateral fields grayish brown with an enclosed series of buffy dots, more distinct posteriorly than anteriorly, the lateral fields set off above by the lateralmost dorsal lines, and below by a series of bluish spots; lower sides spotted with blue. Throat dirtj^ pinkish gray, gular band absent, belly gray. Fore- limbs with scattered pale greenish blue spots, hindlimbs pro- fusely dotted with pale yellow. Tail reddish brown above, grayish blue below, with some blue scales on the sides. Variation: See tables. The large series of A. c. secessa at hand shows little variation in pattern ; the entire animal invariably is quite dull, and the lateral fields are never conspicuous. The longi- tudinal lines do not contrast especially strongly with the dorsal ground color and in some topotypes are very obscure and are seen with some difficulty. In many specimens there are six SCHWARTZ AND KLINIKOWSKI : AMEIVA 469 (rather than seven) dorsal lines, the median line being absent. In- variably the gular band is poorly developed or completely absent ; if the band is present, it does not involve the anterior ventrals but may send some pigment onto the underside of the arms. The coloration of the venter varies from gray and bluish gray to dull orange-gray. There is no sexual dichromatism. Comparisons: A. c. secessa is so dull and drab compared to all other races that no comparison is really necessary. It differs from the spotted races parvoris, ficta, and ahhotti in being lon- gitudinally lined, and from the patternless races hoekeri, richard- thomasi and leheri in having a pattern. It is much duller pat- terned, and likewise differently patterned, from the other lined races — clirysolaema, mnhratilis, alacris, and patterned hoekeri. From jacta and richardthomasi (in the patterned phase), secessa differs in lacking the lateral tigroid markings and in having a quite different dorsal pattern. Perhaps the most cogent compari- son is with clirysolaema which occupies all the adjacent mainland about the Golfe de la Gonave. From clirysolaema, secessa can at once be differentiated by its much more drab coloration and pattern, and by the lack of dotting in combination with lines on the dorsum. Both chrysolacma and secessa are typically 12-row lizards. In fourth toe scales, secessa has a higher mean than any other race, being approached most closely by ahhotti (89.6). In num- ber of femoral pores, secessa is exceeded only by ahhotti, chryso- laema and richardthomasi; in fifteenth verticil scales, secessa, exceeds ficta, richardthomasi, leheri and jacta, and is exceeded by the means of the balance of the subspecies. Remarks: Although A. c. secessa, has presumably evolved from the adjacent A. c. clirysolaema, in dorsal pattern it grossly resembles alacris and procax, but is quite distinct in several features, notably the obscure lateral fields and the lack of a gular band. It seems likely that the nominate race has carried the pattern evolution — i.e., disintegration of the longi- tudinal lines into a series of dots — farther than has the isolated secessa which has become faded and pale in contrast to its more brightly colored neighbor. Another possible origin of secessa is discussed below. 470 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Ameiva chrysolaema DEFENSOR^ iiew subspecies Holotype: MCZ 63379, an adult male, from Mole St. Nicholas, Dept. du Nord Quest, Haiti, one of a series taken 24-29 July 1960 by A. S. Kand and J. D. Lazell, Jr. Paratypes: All from Haiti, Dept. du Nord Quest, as follows: MCZ 63368-72, 63374-78, same data as holotype; MCZ 63364- 67, Jean Rabel, 26 July 1960, A. S. Rand and J. D. Lazell, Jr. ; AMNH 49856-57, Port a I'Ecu, 1 April 1935, W. G. Hassler; USNM 59925, Bale des Moustiques, 3 May 1917, W. L. Abbott ; AMNH 49851-55, river just W Port-de-Paix, 2 April 1935, W. G. Hassler ; MCZ 58014, river just W Port-de-Paix, 2 August 1935, W. G. Hassler. Associated specimens: Haiti, Dept. du Nord Quest: Bombar- dopolis, 1 (MCZ 63381) ; Dept. de VArtibonite, Gros-Morne, 1 (MCZ 63380). Diagnosis: A subspecies of A. c. chrysolaema characterized by a combination of moderate size (males to 126 mm, females to 106 mm snout-vent length), 10 transverse rows of ventrals, low number of fourth toe subdigital scales and scales in the fifteenth caudal verticil, and moderate number of femoral pores ; dorsal pattern a series of six or seven dull longitudinal lines on a tan to brown background, lateral fields dull brown, not especially con- trasting with the dorsal ground color and often with the included light spots in the lateral field much reduced or completely absent, a checkerboard tail pattern (Fig. 8, left), and no indication of a black gular band, black on the anterior ventrals, or extension of black pigment onto the underside of the arms. Distribution: The northwest peninsula of Haiti, from Bom- bardopolis in the south to the vicinity of Port-de-Paix in the northeast, and thence south to Gros-Morne (Fig. 11). Description of type: An adult male with the following meas- urements and counts: snout-vent length 118 mm, tail 242 mm, partially regenerated; ventrals in 38 longitudinal and 12 trans- verse rows; fourth-toe subdigital scales 44 and 44 (total 88), fem- oral pores 19 and 18 (total 37) ; 38 scales in the fifteenth verticil. Dorsal ground color (in preservative) dull brown with a series of seven tan longitudinal lines, the median line the least conspic- uous, all lines disappearing on the neck. Lateral fields brown, with included tan dots only in their posterior thirds, the anterior 1 For the Latin for "defender" in allusion to the English fort at Mole St. Nicholas which guarded the Windward Passage. SCHWARTZ AND KLINIKOWSKI : AMEIVA 471 Fig. 8. Left, Ameiva c. defensor, holotype, MCZ 63379, Mole St. Nicholas, Dept. du Nord Quest, Haiti. Bight, Ameiva c. woodi, unnumbered specimen from MCZ 37583-92, He de la Tortue, Haiti. two-thirds being without dots. Forelimbs grayish tan, vaguely dotted, hindlimbs brown with large pale spots, leaving almost a reticulum of dark brown surrounding the large pale areas. Ven- tral ground color (including throat) bluish gray, no black pig- ment on throat, chest, or undersides of arms. Lower sides with gray and blue markings which are almost tigroid. Tail tan, heav- ily checkerboarded with dark brown above, blue-gray marked with cream below, and with black and some cream on sides. 472 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Variation: See tables. We are somewhat handicapped in discussing A. c. defensor since we have not seen this subspecies in life. Judging from the material at hand, especially the fresh specimens from the Museum of Comparative Zoology (old material in the American Museum from Port-de-Paix and Port a I'Ecu is so discolored that it is completely worth- less insofar as coloration is concerned but does still retain some evidences of pattern), defensor is typically a dull and drab lizard with dorsal coloration of tan to brown with six or seven longitudinal buffy lines. The lateral fields are brown and have the included dots much reduced (often absent an- teriorly) or completely absent. In the latter case the lateral field presents an unbroken brow^n lateral band. In some speci- mens, the longitudinal lines have become more obscure than in the type, due to light pigmented areas in the interline regions, and in one extreme case (IMCZ 63378 — snout-vent 119, and thus not the largest male) the entire back is marbled with dark and light and the sides are tigroid, the latter a condition noted to a slighter degree in some other individuals. No specimen has any indication of black on the chest or undersides of the arms, and the gular black band is lacking completely. The prom- inently checkerboarded tail is a constant feature and is plainly discernible even in old and discolored individuals. Comparisons: A. c. defensor most closely resembles A. c. seccssa from He de la Gonave. However, the reduced or absent dotting in the lateral fields, smaller size, and the generally somewhat brighter dorsal pattern (although defensor is nonetheless a rather drab lizard) will distinguish the two races. Secessa usu- ally has 12 rows of ventrals, defensor usually has 10. From the patternless races, def elisor can be distinguished in having a pattern, and from the spotted subspecies by having a dorsal pat- tern of longitudinal lines. From the other lined races, def elisor differs in the lack of a black lateral field with included yellow dots, and lack of a black gular band. In fourth-toe scales, defensor (82.4) exceeds only ficta (81.9) and jacia (79.7) ; in number of femoral pores, defensor (37.2) exceeds only loekeri (36.5), procax (36.1), umlratiUs (35.6), and alacris (33.8). In fifteenth verticil scales, defensor (38.0) ex- ceeds only leheri (37.9) and jacta (35.7). In counts of fourth- toe scales and fifteenth verticil scales, defensor is quite low in the series of subspecies. SCHWARTZ AND KLINIKOWSKI : AMEIVA 473 Remarks: A. c. defensor is not known to intergrade with any other subspecies ; there are two wide hiatuses, however. The closest approximation of records for defensor (Gros-Morne) and chrysolacma (St. Marc) is 83 kilometers. To the east, there are no specimens available between Port-de-Paix and Cap-Ha'itien, a distance of 70 kilometers. Although A. c. secessa is closer geographically to A. c. chryso- laema, the former is much more similar to defensor than to the nominate race. Such a similarity may be merely convergence and may not reflect direct relationships. It is possible, on the other hand, that defensor has reached the northwest peninsula from GonPive ; it seems hardly likely that the reverse is true — i.e., that Gonave has lieen colonized from the north — consider- ing the proximity of Gonave to the adjacent mainland (21 kilom- eters at its closest point) and its distance from the northwest peninsula (72 kilometers at its closest point). Ameiva chrysolaema woodi Cochran, 1923 Ameiva chrysolaemea woodi Cochran, 1923, Occ. Papers Boston Soc. Nat. Hist., 8:181 (type locality — He Tortue, Haiti). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of large size (males to 141 mm, females to 126 mm snout-vent length), 10 transverse rows of ventrals, low number of fourth toe subdigital scales, moderate number of femoral pores and scales in the fifteenth verticil ; dorsum very dark brown with three to five dull yellowish to buffy stripes or a median dorsal buffy longitudinal band ; the stripes may be variously joined and modified to give rather complex dorsal figures which are derived from the simple five lines (Figs. 8, right; 9) ; sides with vertical tigroid markings, and no black gular band or extensions thereof onto the chest and underside of the arms. Distribution: He de la Tortue, Haiti (Fig. 11). Discmsion: The most strikingly patterned and at the same time most variable of the races is A. c. ivoodi. Basically the dorsal pattern is a series of three to five broad, dull yellowish to buffy longitudinal lines on a very dark brown ground. This basic pattern may be modified in that the area between the two paramedian lines may be filled in with a buffy color so that the back has a lateral pair of pale lines and a middorsal pale zone. From this condition, the balance of the back may be filled in with paler, so that the entire back is marked with a single broad pale middorsal zone. In two individuals, the pale lines have 474 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Fig. 9. Left, Ameiva c. woodi, ASFS X2270, Palmiste, He de la Tortue, Haiti. Eight, Ameiva e. woodi, ASFS X2267, Palmiste, He de la Tortue, Haiti. grossly fragmented and joined randomly, to form a bizarre, longitudinally reticulate pattern -which is quite distinctive. The heads are dull gray-green, with the lores gray. The sides are tan to grayish brown ; this lateral color invades the dark brown back to give a series of alternating dark brown and tan vertical tigroid bars, the tan bars often faintly brick colored at their dorsal points. The throats are dirty pale orange, those of the females usually lighter than those of the males (although one female has a brighter orange throat than any other specimen SCHWARTZ AND KLINIKOWSKI : AMEIVA 475 examined in life). The venter is gray, occasionally with a pale orange wash. No specimen has any indication of a black gular band or anj- black on the chest and undersides of the arms, al- though there may be some isolated black flecking on the chest. The tails are tan dorsall3^ wdth prominent black markings, giv- ing a strong checkerboard effect. The undersides of the tails are gray and often have almost as prominent checkerboarding as the upper surfaces. Variation: See tables. In pattern, no other race of A. chryso- laema is comparable to tvoodi; the three to five longitudinal lines are fewer than in the pattern regularly noted in other subspecies, and the peculiar middorsal broad band, either with or without two dorsolateral lines, and the irregular fragmentation and join- ing of the lines on a dark brown ground are all features which woodi shares with no other race. In lacking a gular band, woodi is comparable only to the adjacent defensor, and seccssa and richardtJiomasi. All other forms have the band present. A. c. ivoodi is a remarkably distinct form; it resembles none of the mainland races and is particularly unlike the adjacent mainland defensor. A. c. ivoodi is a large subspecies, being exceeded only by chrysolaema in size, although procax is equal in snout-vent length. Woodi, in having a low mean (80.5) of fourth-toe scales, exceeds only jaeta (79.7) in this count. In femoral pore count, woodi is exceeded by abhotti, chrysolaema, ricliardthomasi , and secessa, and has a mean femoral pore count equal to those of jacta and leheri. The moderate fifteenth verticil coimt (38.8) of woodi exceeds only richardthomasi, defensor, leheri, and jacta, and is equal to that of ficta. The derivation of woodi must certainly be from the adjacent defensor of the mainland. The lack of a gular band, and black on the chest and undersides of the arms, and the ten rows of ventrals are features in common between the two races. There the resemblances cease, however, since woodi is a boldly and color- fully patterned lizard, whereas defensor is dull and drab. De- fensor likewise does not exhibit any patterns which are reminis- cent of those of vwodi, although the lack of dots in the lateral fields may foreshadow the absence of these fields entirely in woodi. Specimens examined: Haiti, He de la Tortiie, Palmiste. 10 (ASFS X2267-76) ; no specific locality on the island, 29 (MCZ 37583-92 -f 19 unnumbered specimens). 476 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Ameiva chrysolaema regularis Fischer, 1888 Ameiva regularis Fischer, 1888, Jahrb. Wiss. Anst. Hamburg, 5:26 (type locality, Sans Souci, Haiti; this locality is the palace of the same name near Milot, Dept. du Nord, Haiti). Diagnosis: A subspecies of A. chrysolaema characterized by a combination of large size (males to 132 mm, females to 128 mm snout-vent length), usually 12 transverse rows of ventrals, very low number of fourth-toe subcligital scales, moderate number of femoral pores, and high number of scales in the fifteenth verticil ; dorsal pattern a series of five to seven pale yellow lines on a tan to browm ground color, occasionally with a clear tan middorsal zone, neck greenish and dorsal ground color often suffused with Fig. 10. Ameiva c. regularis. ASFS Y-1215, 9 km NW Villa Vasquez, Monte Cristi Prov., Republiea Dominicana. SCHWARTZ AND KLINIKOWSKI : AMEIVA 477 blackish, lateral fields black with an included row of yellow dots (Fig. 10), and black gular band present or absent, when present seldom invading the chest or extending onto the under- side of the arms. Distribution: North central Hispaniola, from Cap-Haitien and Grande Riviere du Nord in the west, east to Fort Liberte, Haiti, and thence to Monte Cristi and throughout the Valle de Cibao as far east as the vicinity of Santiago, Republica Domini- cana; also the Siete Hermanos islands (known from Isla Muer- tos, Toruru, Monte Chico, and Tercero) and Isla Cabras to the north of Monte Cristi (Fig. 11). Discussio7i: Cochran (1941 : pi. 8, figs. B and D) has illustrated two phases in the dorsal pattern of A. c. regularis from Cap- Ha'itien, which is near the type locality of the subspecies. Gen- erally, throughout its wide range on the mainland, regularis is fairly consistent in dorsal pattern. Specimens from the Valle de Cibao, Monte Cristi, and Pepillo Salcedo, which we have seen in life, were brown dorsally, often suffused with blackish, with a greenish wash on the neck, and had five to seven longi- tudinal pale yellow lines. At times there is a clear tan mid- dorsal zone resulting from fusion of stripes and filling in of the interspaces with tan. The lateral field is black and prom- inent with a longitudinal series of yellow dots. The lower sides are dotted with blue-green, and the sides of the belly are bright blue. The ventral ground color varies from grayish to deep dull orange, and the throat from yellowish to gray-orange. The fore- limbs are spotted with blue-green, the hindlimbs with yellow. The gular band may be either present or absent ; if present, it is not extensive and seldom encroaches upon the chest or extends onto the undersides of the arms. There are variants of the above basic pattern, such as that shown by Cochran (1941: pi. 8, fig. D), in which there are faint filigreed lines in a middorsal zone. Some specimens show acces- sory dots between the longitudinal lines, especially posteriorly; in the series from near Villa Vasquez, Monte Cristi Province, two in life clearly showed a secondary dorsal condition, similar to that typical of adult A. c. cJirysolaema, in which the dorsal lines are supplanted by a series of bright yellow dots overlying the fainter longitudinal lines. In general, specimens from near Cap-Haitien seem somewhat darker than those from the xeric Valle de Cibao, but in features of pattern they are not remark- ably different from those from the Valle de Cibao. Occasional 478 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY specimens from the western portion of the range of regularis have the yellow dots in the lateral fields very tiny, and the field thus appears, grossly, to be immaculate, as is characteristic of some defensor. A series of three adult male lizards from Isla Cabras, off the coast just north of Monte Cristi differs from mainland ma- terial in having dark brown lateral fields, and yellow dots on the lower sides. These three specimens also have extensive black gular bands involving the chest and the undersides of the arms. Neither in size nor scalation do there seem to be any differences betw^een these lizards and those from the adjacent mainland. We consider them as regularis since in most features of pattern and coloration they are very close to that race. There are 17 specimens from the Siete Hermanos, a group of seven islets off the mouth of the Kio Yaque del Norte. Of these lizards, ten are from Isla Muertos, two from Isla Tercero, two from Isla Toruru, and three from Isla Monte Chico. In colora- tion, pattern, and scalation they do not differ from mainland specimens, and we regard them as regularis. Variation : See tables. A. c. regularis may be differentiated by its lined pattern from the patternless races — hoekeri, leheri, richardtJiomasi — and those which have distinct patterns (dots, vermiculations, etc.) — parvoris, jacta, ricJiarcWwmasi, fief a, ab- hotti. From the lined races, regularis differs in lacking a dotted and lined dorsum in combination (chrysolaema), in having a dark ground color dorsally and complete and black lateral fields (defensor, secessa), in almost always lacking a pattern of a mid- dorsal tan zone and at times having a black gular band (woodi), and in having twelve rather than ten rows of ventrals (hoekeri, alacris, procax, umhratilis). Begularis most closely resembles alacris and procax; compared with alacris, regularis reaches a larger size and averages fewer fourth-toe subdigital scales (77.8 versus 84.8). From procax, regularis differs in smaller adult size and in often lacking a black gular band, which procax regularly possesses. The two subspecies differ in fourth-toe scales, with regidaris having a lower mean (77.8) than procax (84.8). The north central subspecies is not known to intergrade with either of its neighbors ; there are no specimens to the west between Cap Ha'itien (regularis) and Gros-Morne (defensor) , a distance of 70 kilometers. The closest approximation of regularis (Fort Liberte) and alacris (Cerca-la-Source) is 58 kilometers. The intervening mountains almost certainly completely separate these two subspecies. SCHWARTZ AND KLINIKOWSKI : AMEIVA 479 The distribution of A. c. rcgularis along the northeastern coast of Haiti and thence into the Valle de Cibao in the Republica Dominicana suggests that this form has evolved along the north- ern coast and thence has penetrated into the xeric cul-de-sac of the Valle de Cibao. The affinities of rcgularis with alacris sug- gest strongly that the parent stock has been the latter, yet we cannot visualize any means of dispersal of alacris to the north Haitian coast ; presently at least the Massif du Nord and the Cordillera Central form insurmountable barriers. One route of dispersal suggests itself : the valleys of the Grande Riviere du Nord and the Riviere Bouyaha (the latter a member of the Artibonite system whose upper valleys are occupied by alacris) approach each other in the Departement du Nord. These valleys and their approximation may have offered a means of ingress for Ameiva from the south into the northern Haitian littoral. Specimens examined: Haiti, Dipt, du Nord, Cap-Haitien, 14 (USNM 74075-86, MCZ 37593-94) ; Ti Guinin nr. Cap-Haitien, (not mapped), 24 (UMMZ 122819 [=12 specimens], MCZ 66527- 38) ; Grande Riviere du Nord, 46 (UMMZ 122820 [=12 speci- mens], MCZ 63353-63, 66514-26); Fort Liberte, 6 (USNM 76770-75) ; RepuMica Dominicana, Monte Cristi Prov., Laguna de Salodillo, 7 km S Pepillo Salcedo, 1 (ASFS V1430) ; 4 km E Pepillo Salcedo, 8 (ASFS V1149-55, V1166) ; Isla Cabras, 3 (ASFS V1372-74) ; Monte Cristi, 1 (MCZ 58018) ; 2 km SE Monte Cristi, 5 (ASFS V1210-12, V1284-86) ; 9 km NW Villa Vasquez, 14 (ASFS V1214-25, RT 811-12) ; 5 km W Guayubm, 15 (ASFS V1494-508) ; 7 km N Guayubin, 13 (ASFS V1471-83) ; Valverde Prov., 9 km N Los Quemados, 1 (ASFS V1766) ; 7 km E Valverde, 10 (ASFS V2931-40) ; 2 km E Esperanza, 5 (ASFS V1755-59) ; Santiago Prov., 7 km W Santiago, 2 (ASFS V2925- 26) ; Santiago and vicinity, 6 (MCZ 58665-66, 58668-71) ; Siete Hermanos, Isla Muertos, 10 (ASFS V1590-95, RT 826, USNM 76733-35) ; Isla Monte Chico, 3 (USNM 76715-17) ; Isla Ter- cero, 2 (USNM 76736-37) ; Isla Toruru, 2 (ASFS V1573-74). DISCUSSION Before proceeding to a discussion of the variation and possible history of Ameiva chrysolaema in Hispaniola, we would like to bring out several facts which seem especially worthy of mention. The distribution of patternless races, or at least races which have some patternless members (leheri, hoekeri, richardthomasi), 480 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY SCHWARTZ AND KLINIKOWSKI : AMEIVA 481 is especially interesting. Of these subspecies, one (leheri) is iso- lated on the south shore of His^Daniola below the La Selle-Baoruco massifs; leheri approaches, insofar as known without intergrada- tion, the very different patterned race ficta on the Peninsula de Barahona. Another (hoekeri) has patterned individuals, and is surrounded by three patterned races, of which it intergrades with one {alacris), is separated from another (umhratilis) by the Rio Yaque del Sur, and from the third {procax) by a distinct and dramatic change of environment. Finally, richard- fhomasi on Saona has patterned and patternless individuals; this subspecies is related most closely to jacta. Three races are dorsally spotted : ahhotti and ficta on Beata and the Peninsula de Barahona, respectively, and parvoris on the southeastern coast of Hispaniola. Parvoris is separated from its neighbor to the west, procax, by the Rio Ozama, and from its eastern neighbor, jacta, by a wide gap which is apparently })resently uninhabited by A. chrjjsolaema. The occurrence of parvoris on Isla Catalina, to the east of the known mainland distribution of that race, is noteworthy. Of the striped races, chrysolaema stands alone in its large size and its style of dorsal patterning, a combination of lines and dots. Chrysolaema is known to intergrade with alacris, another striped race, in the vicinity of Mirebalais, but no intergrades are known between chrysolaema and unihratilis in the Cul de Sac- Valle de Neiba. rmhraiilis resembles the striped phase of hoekeri to some extent ; the two are not known to intergrade. The races procax, ala<;ris, and regularis are all comparably striped ; of them, alacris and procax have 10 rows of ventrals, whereas regularis has 12 rows of ventrals. The range 0/ alacris is sep- arated from that of procax by the intervening and quite different hoekeri. Defensor, by virtue of its pallid coloration and drab pat- tern, stands alone among the mainland races, but it is approached somewhat by the drab secessa from Gonave ; secessa has 12 rows of ventrals. rkfensor 10. The Tortue subspecies woodi is very distinctive, I)ut logically must have been derived from either defensor or regularis, the only two races on the north coast. The Sierra de Xeiba and the Montagues du Trou d'Eau form the northern boundary of the Cul de Sac-Valle de Neiba plain, which is in actuality a fossil strait that once separated His- paniola into two distinct islands, the north and south islands The south side of the same plain is bounded by the Massif de la Selle and its associated northern ranges (Morne I'Hopital, ]\Iont 482 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY des Enfants Perdus) and the Sierra de Baoruco. Considering only this region, one is struck by the amazing diversity of the four races of A. chrysolaema associated with it : chrysolaema in the northwest, umhratilis in the northeast, ficta in the southeast and leberi in the southwest. Here we are involved with four races whose patterns are radically different — chrysolaema dark with longitudinal lines and dots, umbratilis pale with lines, ficta with large dorsal spots, and leheri without pattern. It has become customary to catalogue, if possible, Hispaniolan reptiles and amphibians into either north or south island species — i.e., depending upon their present and presumed past distribu- tion. Although this is somewhat difficult in the present case, we feel that A. chrysolaema is a north island species. Except for the isolated occurrence of A. c. chrysolaema at Aquin on the Tiburon Peninsula (based on a single specimen), this south- western extremity of Hispaniola lacks the species; the otherwise westernmost record is from Pere, near Leogane. The occurrence of the very different race ficta on the Peninsula de Barahona (and its relative ahhotti on Beata), as well as leheri to the west along the south coast, indicates that the A. chrysolaema stock was long isolated on the southeastern portion of the south island, where ficta evolved in isolation from the northern mass of the species. The presence of leheri along this south shore is most puzzling, especially since it is geographically closest to ficta (see Remarks under A. c. ficta for additional comments), and since it also resembles hoekeri far to the northeast. Perhaps leheri is the more ancient of the two mainland south island races, and its present rather restricted distribution a mere remnant of a range which was once more extensive, especially to the west toward Jacmel. If such is the case, leheri might be regarded as a sub- species derived from (pre) chrysolaema; a possible source of colonization from the northern shore of the Tiburon Peninsula might be the Vallee de Trouin, the low pass between the north and south shores of the peninsula. It is of course quite possible that additional collecting in the vicinity of Jacmel or between that city and Saltrou will reveal the presence of leheri; it is also possible that chrysolaema or chrysolaema X leheri inter- grades may l)e found in the Vallee de Trouin as well. On the north island, and including the Cul de Sac-Valle de Neiba, we visualize the old coast inhabited by two races, chryso- laema and umhrotilis, much as today procax and parvoris occur along the south shore of the eastern Republica Dominicana. With SCHWARTZ AND KLINIKOWSKI : AMEIVA -483 the closure of the strait, each of these races has expanded into the resulting xeric plain, although to the northwest along the Golfe de la Gonave, chrysolaema still occupies the narrow coastal plain and adjacent xeric foothills much as it may formerly have done farther south. Once across the plain, chrysolaema has ex- tended its range inland to some extent (Petionville), and to the west (Pere, Momance). Vmhratilis, on the other hand, has not been able to penetrate far into the adjacent mountains, although it does reach an elevation of 1000 feet near El Naranjo. The range of nnihratilis is bounded on the north by the valley of the Rio Yaque del Sur. To the east of the Rio Yaque del Sur are a series of four, more ©r less coastal, races. Of these, hoekeri, immediately to the north and east of the Yaque, is rather like umbratilis in its patterned phase. We consider hockcri as a direct derivative of umhratilis and restricted to the Llanos de Azua. The next three races — procax, parvoris, jacta — show increasingly scattered patterns of distribution to the east, with jacta apparently the most iso- lated. As noted previously, procax and hoekeri approach one another in the vicinity of Bani, precisely in the area of rapid transition from the xeric Llanos de Azua to the more mesic coastal areas to the east. The Rio Ozama separates procax from parvoris, which is known from only two localities on the main- land and from a slightly differentiated population on Isla Cata- lina. We consider hoekeri, procax, and parvoris as a more or less sequential coastal series still maintaining its integrity in response to environmental and geographical influences. Jacta, on the other hand, along with richardthomasi, represents a very different sort of lizard. We feel that the jacta-richard- thomasi populations at one time (and perhaps still) occupied most of the extreme eastern end of the island. The presently restricted and scattered distribution and records for not only jacta but also parvoris, as well as the isolated occurrence of parvoris on Catalina, suggest strongly that the range of A. chrysolaema in this region is retracting, leaving isolated outliers which may be indicative of former populations. The absence of records of the species between San Pedro de Macoris and Juan- illo, as well as only two general localities of parvoris on the mainland, add substance to this supposition. The eastern dis- tribution of Ameiva lineolata tends also to bear out this conten- tion. 484 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Of the remaining- races from the southern part of the north island, only the strii)ed alacris is left. Alacris is so like procax in many features that it is difficult not to associate the two. On the other hand, they are separated presently by hoekeri, with which race alacris intergrades, as it does with chrijsolaema on the west. Tt is possible that alacris and procax were at one time confluent, but that bockcri has pushed between them, thereby severing- any direct genetic continuity between the two. An- other possibility is that alacris has been derived from chryso- laema, either across the Montagues de Trou d'Eau or along the valley of the Riviere de I'Artibonite. Continued evidence of intergradation between these two races near ]Mirebalais lends support to this possibility. A. c. defensor on the northwestern peninsula has obviously been long isolated from its more southern relatives. Presumably it has been derived from chrysolaema. The Gonave race secessa resembles defensor in pattern and cohn-ation, and it is possible that Gonave was colonized from tlie north (defensor) rather than from the adjacent mainland [chrysolaema) . The latter, however, seems more likely both on the basis of proximity and what is presently known of the origin of the Gonave herpetofauna. The relationships of regularis seem closest to the procax-alacris pair; possibility of origin of regularis from alacris via the Arti- bonite system and thence to the Grande Riviere du Xord has already been discussed. Other possibilities are an old origin from defensor (although this is not particularly appealing) or an origin from procax via the central valley to the east of the Cordillera Central in the Republica Dominicana. The only evi- dence against this is that A. chrysolaema does not occupy this val- ley today, although procax occurs at its southern end and regularis at its northern end uear Santiago. Finally, A. c. woodi on Tortue, although closer geographically to defensor than to regularis, seems closer in some characteristics to regularis than to defensor. There are, however, tendencies of pattern in defensor which herald the extreme peculiarities of woodi patterns. On the other hand, occasional regularis have dorsal patterns like some woodi. It seems more likely that ivoodi is a direct derivative of defensor. SCHWARTZ AND KLINIKOWSKI : AMEIVA 485 LITERATURE CITED Barbour, Thomas, and G. Kingley Noble 1915. A revision of the lizards of the genus Ameiva. Bull. Mus. Comp. Zool., 59(6):417-479. Cochran, Doris M. 1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: i-vii + 1-398, 12 pis., 120 figs. Maerz, a., and M. Rea Paul 1950. A dictionary of color. New York, McGraw-Hill Book Co., pp. i-vii, 1-23, 137-208, 56 pis. Mertens, Robert 1938. Aniphiliien und Reptilien aus Santo Domingo, gesanimelt von Dr. H. Boker. Senckenbergiana, 20: 332-42, 6 pis. 1939. Herpetologische Ergebnisse einer Reise naeh der Insel Hispan- iola, Westindien. Abh. senckenberg. naturf. Ges., 449: 1-84, 10 pis. Schwartz, Albert (in press) The Ameiva (Reptilia, Teiidae) of Hispaniola. I. Ameiva lineolata Dumeril and Bibron. Caribbean Jour. Sci. (Received February 25, 1965.) 486 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Subspecies chrysolaema woodi procax richardthomasi parvoris secessa jacta regularis umbratilis boeheri alacris defensor ficta abbotti leberi 160 141 141 137 137 135 134 132 130 126 126 126 121 117 111 130 126 116 124 113 111 128 112 111 109 106 113 108 104 Table 1. Subspecies of Ameiva chrysolaema ranked according to snout- vent length (in mm) of largest male for each race. Transverse ventrals Table 2. Subspecies of Ameiva chrysolaema ranked according to means of number of fourth-toe subdigital scales; each race is also characterized by the modal number of transverse rows of ventral scales (but see discussions of umbratilis, procax, and richardthomasi). N ^ number of specimens ex- amined. SCHWARTZ AND KLINIKOWSKI : AMEIVA 487 Subspecies abbotti chrysolaema richardtJiomasi seoessa jacta leberi woodi ficta parvoris regularis defensor boeTceri procax umbratilis alacris Table 3. Subspecies of Ameiva chrysolaema ranked according to mean number of femoral pores; N = same as in Table 2. Scales in 15th caudal verticil Subspecies Mean and extremes chrysolaema alacris regularis procax boeTceri umbratilis parvoris abbotti secessa ficta woodi richardthomasi defensor leberi jacta Table 4. Subspecies of Ameiva chrysolaema ranked according to mean num- ber of scales in fifteenth caudal verticil ; N = same as in Table 2.