Fh aye VAG YR HOD ya) ats pat 4 ) Bike. : oN a Ae hye \ oR iaate 1 ba Say ss Ay 3 bate “Bet M44 utes , ee 4g UAE Se Daca Teka inti ate SAM SST AF See wae Wee nea a eee AWN + aS A mA a —— en be OP DEC 17 1971 BHARV ARO UNiversiTs POSTILLA PEABODY MUSEUM YALE UNIVERSITY NUMBER 154 1 DECEMBER 1971 LEPOSOMA PERCARINATUM, A UNISEXUAL SPECIES RELATED TO L. GUIANENSE; AND LEPOSOMA IOANNA, A NEW SPECIES FROM PACIFIC COASTAL COLOMBIA (SAURIA, TEIIDAE) THOMAS UZZELL JOHN C. BARRY POSTILLA Published by the Peabody Museum of Natural History, Yale University Postilla includes results of original research on systematic, evolution- ary, morphological, and ecological biology, including paleontology. Syntheses and other theoretical papers based on research are also welcomed. Postilla is intended primarily for papers by the staff of the Peabody Museum or on research using material in this Museum. Editors: Zelda Edelson, Elizabeth G. Weinman, Elise K. Kenney. Postilla is published at frequent but irregular intervals. Manuscripts, orders for publications, and all correspondence concerning publications should be directed to: Publications Office Peabody Museum of Natural History New Haven, Conn., 06520, U.S.A. Lists of the publications of the Museum are available from the above office. These include Postilla, Bulletin, Discovery, and special publica- tions. Postilla and the Bulletin are available in exchange for relevant publications of other scientific institutions anywhere in the world. Inquiries regarding back numbers of the discontinued journal, Bulletin of the Bingham Oceanographic Collection, should be directed to: Walter J. Johnson, Inc. 111 Fifth Avenue New York, N.Y. 10003. LEPOSOMA PERCARINATUM, A UNISEXUAL SPECIES RELATED TO L. GUIANENSE; AND LEPOSOMA IOANNA, A NEW SPECIES FROM PACIFIC COASTAL COLOMBIA (SAURIA, TEIIDAE) THOMAS UZZELL Department of Biology and Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06520 JOHN C. BARRY Department of Geology and Geophysics, Yale University, New Haven, Connecticut 06520 (Received May 19, 1971) ABSTRACT Two distinct populations of Leposoma occur in the Guianan highlands area of northern South America. Leposoma percarinatum Miller is distinguished from L. guianense Ruibal by a higher number of transverse rows of dorsal scales (36-39, mean 37.5 in L. percarinatum; 31-35, mean 32.6 in L. guianense) as well as by more transverse rows of ventral scales and more scales around the midbody region. In L. percarinatum, the third and especially the fourth pairs of chin shields are much reduced compared to the conditions in L. guianense; the scale behind the longest supralabial is as small as the tem- poral scales, rather than larger as in L. guianense. L. percarinatum usually has a light line from the hindlimb insertion along the tail; this is absent in L. guianense. The interparietal scale in L. percarinatum is much larger than in L. guianense, but the head is relatively narrower. The distinctive long middle preanal scale noted by Ruibal is not a constant feature of L. guianense. All 30 specimens of L. percarinatum are female. We have examined 17 specimens of L. guianense; seven are male, 10, female. L. percarinatum is probably a unisexual taxon. Male specimens reported are probably L. guianense. In coloration, L. percarinatum seems to be intermediate between L. guianense and L. parietale, a species found on the Amazonian slopes of Peru, Ecuador and Colombia. In scale counts, size of fourth pair of chin shields, and head width, L. parietale and L. guianense are alike. The interparietals of L. percarinatum and L. parietale are both large. Geographically and morpho- logically, it seems likely that L. parietale and L. guianense hybridized to give rise to L. percarinatum. If this is so, these bisexual forms are probably distinct species, although the distributions and general similarities suggest that they represent subspecies of a single species. Leposoma ioanna, new species, is closely related to L. parietale. The two known specimens come from near Buenaventura, Valle del Cauca, on the Pacific coast of Colombia. L. ioanna is distinguished from L. parietale by having the third pair of chin shields separated from the infralabials by two small scales, and by a much reduced fourth pair. It may have a larger body size and lower mean numbers of dorsal scale rows, transverse ventral scale rows, and scales around midbody region. Two Chocéan specimens reported previously as Alopoglossus copii are Leposoma southi. Both L. ioanna and L. southi are species of Leposoma from the Pacific coast of Colombia, whence no species has been reported previously. L. southi also occurs in Costa Rica and Panama east as well as west of the Canal Zone. Hemipenes of Leposoma have two characteristic rows of calcareous spines in each half. Features of the hemipenis of L. parietale are also seen in the hemipenis of Arthrosaura kockii, supporting the relation of the genera sug- gested on other bases. Evidence of past hybridization of L. parietale with L. guianense (the uni- sexual L. percarinatum) and the disjunct L. ioanna, which is related to L. parietale, indicate considerable changing of distributions of the genus Lepo- soma. Dates of changes in distribution are unknown. POSTILLA 154: 39p. 1 DECEMBER 1971 CONTENTS ABSTRACT I Introduction II Leposoma percarinatum, a unisexual species related to Leposoma guia- nense and Leposoma parietale By Thomas Uzzell and John C. Barry III Leposoma ioanna, a new species from the Pacific coast of Colombia, and records for Leposoma southi, new to the fauna of Colombia IV A key for identifying lizards of the genus Leposoma V The hemipenis of species of Leposoma The senior author alone is responsible for the abstract and sections I, III, IV, and V. 4 POSTILLA 154 I. INTRODUCTION The small teiid lizards of the genus Leposoma, a member of Boulenger’s (1885) group II of the family Teiidae, were reviewed by Ruibal in 1952. Ruibal examined 19 females but no males of L. percarinatum, and 2 females but no males of Leposoma guianense, a taxon that he described as new. Re- cently, considerable material of these two taxa was assembled to investigate their relations to each other and to L. parietale. The material includes all that Ruibal saw, and some additional specimens. The additional material consists of 11 additional specimens of L. percarinatum (all females), and 15 additional specimens of L. guianense (7 males and 8 females). Although the feature of the preanal scales described as diagnostic for L. guianense does not distinguish the taxon, several additional characters, in- cluding features of the interparietal mentioned by Ruibal, do distinguish it. Among the specimens examined are 11 from the Pacific lowlands of Co- lombia, whence Leposoma has not previously been recorded. Two of these, while rather similar to L. parietale of the Amazonian slopes of the Andes in Ecuador, Colombia and Peru, show certain distinctive features. Because there probably has been no recent contact between the lowland Pacific coast popu- lation and the lowland Amazonian population, the coast form is described as a distinct species. The other specimens are L. southi, heretofore known only from Costa Rica and adjacent Panama east into the Canal Zone. ABBREVIATIONS AMNH American Museum of Natural History, New York ANSP Academy of Natural Sciences of Philadelphia BMNH British Museum (Natural History), London FMNH Field Museum of Natural History, Chicago MCZ Museum of Comparative Zoology, Harvard University MLS Museo de La Salle, Bogota MNHN Muséum National d’Histoire Naturelle, Paris SMF Senckenberg Museum, Frankfurt-am-Main UKMNH University of Kansas Museum of Natural History USNM United States National Museum UMMZ University of Michigan Museum of Zoology II]. Leposoma percarinatum, A UNISEXUAL SPECIES RELATED TO Leposoma guianense! By Thomas Uzzell and John C. Barry Unisexual species of lizards have been reported on many occasions since Darevsky and Kulikova (1961), Maslin (1962), Minton (1958), Tinkle (1959), and Zweifel (1965) first drew attention to them. At present more than 20 different unisexual taxa of lizards are known with some degree of assurance, 1. Our study parallels one by Marinus S. Hoogmoed, Rijksmuseum van Natuurlijke Historie, Leiden, who has independently come to essentially the same conclusions. LIZARDS OF THE GENUS LEPOSOMA 5 and others doubtless will be reported as additional refined variational studies of lizards are undertaken. The unusual sex ratio of Leposoma percarinatum was noted by Ruibal in 1952 when unisexual vertebrates were almost unknown. The closely related L. guianense Ruibal (1952) was only known from 2 females. We have ex- amined additional material of both taxa and we can demonstrate that in the Guianan highlands region of South America (Venezuela, the Guianas, and adjacent Brazil and Colombia) there exist 2 separable populations of fe- males. One of these (L. percarinatum) is known only from females. Both males and females are known for the other (L. guianense). Leposoma guianense and Leposoma percarinatum may be distinguished by the following characteristics. 1. Leposoma guianense has relatively fewer body scales than L. percari- natum. Table 1 summarizes data on the number of transverse rows of ventral TABLE 1. MAXIMUM LENGTHS AND RANGES AND MEANS FOR THREE SCALE COUNTS IN FIVE SPECIES OF Leposoma Maximum Dorsal Transverse Scales body length scale rows of around (mm) rows ventrals midbody L. guianense 762 35 32-34 22-24 23-25 (32.9) (23.0) (24.0) 109 9 38 31-35 21-23 23-26 (32.4) (2235) (24.4) L. percarinatum 302 9 35 36-39 24-27 23-28 (37.5) (25.5) (25.4) L. parietale1 29-35 20-24 21-25 2b So 36 (32.9) (22.4) (23.6) 2292 9 39 31-36 2124 22-26 (33.2) (22.4) (23.7) L. ioanna PREM) 41 28, 30 20521 21, 22 L. southi 34 6 40 27-29 19-20 21-23 (2ilev) (19.7) (21.7) 62° 37 28-29 20-21 21-22 (28.5) (20.3) (23) 1Based on AMNH _ 56260-62, 106635-37, ANSP 25507-08, FMNH 45477, MCZ 61156, UKMNH 98601-03, 121784-85, 121789-97, 121799-803, 121807-08, 121811-26. 6 POSTILLA 154 scales between the collar and the preanal scales, on scale rows around the midbody region, and on the number of dorsal scale rows between inter- parietal and posterior margin of hind limbs. Each of these characteristics alone suffices to identify most of the specimens examined. 2. Leposoma guianense, as noted by Ruibal (1952), has a very large inter- parietal. The length of the interparietal, of each parietal, and the width of the interparietal at its anterior and posterior corners were measured (Fig. 1). \ woes Anterior interparietal Parietdl--+-.....f el. width length Interparietal-"""" | length **tesseseces POSTEFIOF ; interparietal width FIG. 1. Sketch of head scales of Leposoma, showing dimensions measured. The relative lengths of the parietal and interparietal are compared in Figure 2. The area of the interparietal was estimated as the mean of anterior and pos- terior widths times the length. The area of the interparietal changes with body length. Interparietal area relative to body length for L. guianense and L. percarinatum is compared in Figure 3. The number of transverse rows of dorsal scales and the relative interparietal area distinguish the same individuals (Fig. 4). 3. Both Leposoma guianense and L. percarinatum usually have 6 supra- labials. In both, the last supralabial is the longest. Following the longest supra- labial in L. guianense, there is a scale on the lip line that is larger than the LIZARDS OF THE GENUS LEPOSOMA 7 1.9 E e = = 1.7 °° ‘ $ 2 Oo. WwW 88 é 4 1.5 2 ° : a = O 8 e ° = 3 8 0 is «= 5 a oO og e é 7 On Oe q ° | = e ee /eposoma guianense = 09 © Leposoma percarinatum Se hood ole Mel co eo ei eee .o INTERPARIETAL LENGTH (mm) FIG. 2. Mean parietal length as a function of interparietal length in Leposoma guianense and L. percarinatum. adjacent temporal scales (Fig. 5); all of the specimens separated by number of dorsal scale rows and relative interparietal area are also separated by this character. 4. Both Leposoma guianense and L. percarinatum have a median unpaired chin shield followed by two pairs of chin shields that are in contact with each other medially and with the infralabials laterally. Both also have a third pair of chin shields that are separated on the midline, and that are separated from the infralabials by a single small scale (Fig. 6). In L. guianense, the third pair of scales is followed by a fourth, smaller pair of chin shields, separated from each other by a scale and from the infralabials by 3 to 4 scales counted along the posterior margin of the third pair of chin shields. In L. percarinatum, the largest scale behind the third pair of chin shields (and indeed, the third pair of chin shields themselves) is considerably reduced (Fig. 6). All specimens distinguished by the preceding characters are also distinguished by this char- acter. 5. Specimens of Leposoma percarinatum have relatively narrower heads than specimens of L. guianense (Fig. 6). 6. Specimens of Leposoma percarinatum usually have a distinct light line along the side of the tail posterior to the insertion of the hind limb. This line is usually absent in L. guianense. The suite of characters above show a high degree of concordance. Several of the characters (numbers of dorsal and ventral scales, number of scales around the midbody region, approximate interparietal area relative to body 8 POSTILLA 154 length, mean parietal length relative to interparietal length, and parietal width relative to body length) were used in a canonical analysis (BMDO7M; Dixon, 1968). The distribution of individuals of L. guianense and L. percarinatum along the first and second canonical axes confirms the morphological distinct- ness of these two forms (Fig. 7). The specimens examined come from generally sympatric areas (Map 1). Specimens have been collected together at few localities, but in Guyana, for instance, where both species are known, there is no greater similarity than elsewhere. For this reason, we believe that the above characters delimit two distinct biological populations. Of the specimens examined, all assigned to L. percarinatum are females, whereas 7 out of 17 assigned to L. guianense are males. Since even moderate samples of other species of the genus Leposoma contain both males and fe- males, we feel that their absence in the known sample of L. percarinatum (30 es) e¢ / eposoma guianense ¢ 7.0 o Leposoma percarinatum e 6.5 6.0 I8 20 22 24 26 28 30 32 34 36 (38045 SNOUT-VENT LENGTH (mm) FIG. 3. Approximate interparietal area as a function of body length in Leposoma guianense and L. percarinatum. LIZARDS OF THE GENUS LEPOSOMA 2 specimens) is strong evidence that males do not occur in this taxon. This be- lief is reinforced by the numerous individual collections that have been drawn together to form the known sample. The following general taxonomic accounts describe the two Guianan species. Leposoma percarinatum (MULLER) Hylosaurus percarinatus Miiller, 1923, Zool. Anz. 57: 146. Leposoma taeniata Noble, 1923, Zoologica 3: 303. Hylosaurus muelleri Mertens, 1925, Senckenbergiana 7: 76. DESCRIPTION. Head narrow; dorsal head scales longitudinally striate. Fronto- nasal single, wider than long, separated from anterior supraoculars. Nasal divided, elongate posteriorly. Loreal divided: lower segment small. Prefrontals small. Frontal elongate, hexagonal. Frontoparietals short, joined by a rela- tively long suture. Interparietal large, hexagonal, about as wide as long. 39 Oo 6.6 e6 / eposoma guianense © Leposoma percarinatum 38 ©0@m@ 8 00 oT om Oe Of 36 fore o©) 55 e 34 ve é DORSAL SCALE ROWS 0.08 0.10 0.12 0.14 0.16 0.18 0.20 0.22 INTERPARIETAL AREA (mm®) SNOUT-VENT LENGTH (mm) FIG. 4. Number of dorsal scales along midline between occiput and hind limb margin as a function of interparietal area relative to body length for Leposoma guianense and L. percarinatum. 10 POSTILLA 154 Parietals smaller and shorter than interparietal; anteriormost part of each parietal usually projecting farther forward than anteriormost part of the interparietal. Supraoculars 4; superciliary series complete; suboculars 4-6. Temporals small, keeled. Upper labials 6, the last longest; next posterior scale as small as adjacent temporals. A mental followed by one unpaired and three paired chin shields; two anterior pairs in contact medially, third pair sep- arated by a small scale; two anterior pairs in contact with lower labials; third FIG. 5. Lateral view of the heads of Leposoma guianense and L. percarinatum (x10). Above, UMMZ 46770, adult female, holotype of L. guianense; below, AMNH 21266, adult female, holotype of L. taeniata. In L. guianense, the scale along the lip line behind the longest supralabial is markedly larger than the adjacent temporal scales; in L. percarinatum, this scale is nearly coequal in size with the adjacent temporal scales. LIZARDS OF THE GENUS LEPOSOMA i FIG. 6. Ventral view of the heads of Leposoma guianense and L. percarinatum (x10). Above, UMMZ 46770, holotype of L. guianense; below. AMNH 21266, holotype of L. taeniata. In L. guianense, the third and especially the fourth pair of chin shields are considerably larger than in L. percarinatum. The head is relatively wider in L. guianense than in L. percarinatum. pair separated from lower labials by a single small scale. Pregulars (Ruibal, 1952) flat, of variable sizes and shapes. A granular gular crease. Gulars in 9-10 transverse rows. Ventrals imbricate, in longitudinal and 24-27 (mean 25.5) transverse rows between collar and preanals. Dorsals leaf-shaped, keeled, imbricate, in 36-39 (mean 37.5) trans- verse rows. Caudals in longitudinal and transverse rows. Six preanal scales; two on midline, of which one is anterior and the other marginal, these flanked by two large marginal scales, which in turn are flanked by two smaller marginal scales. Adult females usually with 2 preanal pores; no femoral pores. i POSTILLA 154 é€e Leposoma guianense c'o Leposoma porietale w Leposoma percarinatum 4 Leposoma /oanna + Mean SECOND CANONICAL VARIABLE -7.8 -5.8 -38 -1.8 0.2 oe 4.2 6.2 8.2 FIRST CANONICAL VARIABLE FIG. 7. Distributions and means for individuals of four species along the first and second canonical variates. The characters on which the analysis was based include the number of dorsal and ventral scales, the number of scales around the midbody region, the length of parietals relative to interparietal length, the approximate area of the interparietal relative to body length, and the width of the interparietal relative to body length. The first canonical variate (x axis) represents primarily number of body scales; the second (y axis) primarily features of the interparietal. Together, these two canonical variates account for more than 95% of the total variance. Ranges for males and females of the two species represented by both sexes over- lap considerably, and the means are quite close. L. percarinatum is quite distinct, on both axes, from L. guianense and L. parietale. The two specimens of Leposoma ioanna, new species, are most like specimens of L. parietale, but rather markedly set off from them. Body color varies from light brown to reddish brown. Two thin, irregular, dark dorsolateral bands; these markings fade with age. The markings are most conspicuous anteriorly. REMARKS. The holotype of Hylosaurus percarinatus was collected by Lorenz Miller, probably in 1910, in Peixe Boi, Para, Brazil (Map 1). It was cata- logued as 140/1911 in the Zoologische Staatssammlung, Munich. Uzzell was unable to locate the specimen in Munich in 1968; it probably was destroyed during World War II. A question mark in Miiller’s original description may refer to maturity or to sex. Miller reported that the type was 34 mm snout-to- vent, and that it lacked femoral pores. Since in L. guianense, males 34 mm LIZARDS OF THE GENUS LEPOSOMA 15 Sy—|@) L. quionense MAP 1. The Guianan region of northern South America, showing localities for specimens of Leposoma guianense and L. percarinatum. Certain localities discussed in the text are indicated. snout-to-vent have striking femoral and preanal pores, the specimen appar- ently was a female. At 34 mm body length, it was certainly mature, since specimens of this body length often contain oviducal eggs, one per oviduct. Characteristics of the holotype of L. percarinatum that identify it as a member of the taxon to which we apply the name include the number of dorsal scale rows, the number of scales around the midbody area, and the number of ventral scale rows (Table 2). The large scales behind the third pair of postmentals and behind the longest supralabial, characteristics of L. guianense, possibly would have been mentioned by Miiller had they occurred. The sex is appropriate for the present assignment. We have not examined specimens of this taxon from Para. The holotype (AMNH 21266) and three paratypes (UMMZ 58420; AMNH 21263, 21268) of Leposoma taeniata each show the distinctive fea- tures we ascribe to L. percarinatum. All four specimens of the type series examined are from Kartabo, Guyana. Three other paratypes, one at the British Museum (Natural History) and two at the Museum of Comparative 14 POSTILLA 154 TABLE 2. CHARACTERISTICS OF SEVERAL HOLOTYPES IN THE GENUS Leposoma BS ie, ie, jes je. jis natum taeniatal muelleri guianense1 parietale ioanna Sex 2 2 on 2 3 2 Snout-vent length (mm) 34 35) 30 36 32 41 Dorsal scale rows 36 38 391 31 — 28 Transverse rows of ventrals 25 26 25 21 — 20 Scales around mid- body region 26 24 Poy 23 — 21 1Our data. Zoology, have not been examined. Noble (1923) mentioned 8 paratypes, but we have traced only 7. Although the holotype of Hylosaurus muelleri (SMF 11754, formerly 5385a) was described as possibly an adult male, femoral and preanal pores are absent; the specimen is an adult female (Uzzell, personal observation, 1968). The scale counts (Table 2) indicate that the specimen is L. percari- natum. The distinctive characteristics of the scale behind the longest upper labial and behind the third pair of postmentals have been compared for us with photographs of the holotypes of L. guianense and L. taeniata by Konrad Klemmer. The holotype of L. muelleri resembles L. taeniata in both features. The type locality given by Mertens (1925) as Inirida, southern Venezuela. In the 1890s southern Venezuela extended west beyond the Rio Orinoco to about 69° W. It thus would include the lower reaches of the Rio Inirida, presently in Colombia (Map 1). This is apparently the area from which the specimen came (Burt and Burt, 1931; Ruibal, 1952). Da Cunha (1961) reported 35 specimens including 18 males of L. percari- natum, all from the upper Rio Maraca, Amapa, Brazil (Map 1). We have seen none of these specimens. The close similarity of L. guianense and L. percarinatum makes us believe that da Cunha’s series included predominantly or entirely L. guianense, rather than L. percarinatum. Da Cunha reported that his specimens had 33 to 38 transverse rows of dorsal scales, and averaged 27 scales around the midbody region. These values suggest a mixture of specimens, but may be due in part to different ways of counting the scales. SPECIMENS EXAMINED. Brazil: Amazonas: near Salto da Hua (100-200 m) USNM 83573. Guyana: Demerara: Dunoon (0-100 m) UMMZ 46769; near MacKenzie (0-100 m) BMNH 1934.11.1.102; Wismar (0-100 m) AMNH 58981; Essequibo: Aishalton (Ishertun; 100-200 m) AMNH 61418 (3 speci- mens), 61419 (4 specimens); 15—20 miles above Bartica, upper Cuyuni River, Camp 1 (0-100 m) BMNH 1934.11.103; Chenaponu River (450 m) BMNH LIZARDS OF THE GENUS LEPOSOMA 15 1905.11.1.6; Kamakusa (0-100 m) AMNH 25082-83; Kartabo (0-100 m) AMNH 21266 (holotype of Leposoma taeniata), AMNH 21263, 21268; UMMZ 58420 (paratypes of Leposoma taeniata), AMNH 46433-35; Kuyu- wini Landing (200 m) AMNH 57455; Potaro (0-100 m) BMNH 1970.720-21: Tukeit Falls (Tuheit Hills; 0-100 m) UMMZ 63050; Winiperu (0-100 m) BMNH 1968.878. Venezuela: Amazonas: Camp La Culebra (200-500 m) MCZ 58347, 101884; 12 miles N of Esmeraldas (100-200 m) AMNH 39320; Rio Pescada, Mt. Duida (100-200 m) AMNH 36637. Leposoma guianense Ruibal Leposoma guianense Ruibal, 1952, Bull. Mus. Comp. Zool. 106: 489. DESCRIPTION. Head broad; dorsal head scales longitudinally striate. Fronto- nasal single, wider than long, separated from anterior supraoculars. Nasal usually undivided, only slightly elongate posteriorly. Loreal divided; lower segment large. Prefrontals shorter than frontonasal. Frontal long, hexagonal. Frontoparietals elongate back along posterior supraoculars, joined by a rela- tively short suture. Interparietal long and wide, hexagonal. Parietals smaller, barely more than half as long as interparietal; anteriormost part of inter- parietal projecting farther forward than anteriormost part of each parietal. Supraoculars 4; superciliary series complete; suboculars 4-6. Temporals small, keeled. Upper labials 6, the last the longest; an enlarged scale following the longest supralabial. Lower labials 5—6, the third the longest. A mental fol- lowed by one unpaired and three paired chin shields; two anterior pairs in contact medially, posterior pair separated by a small flat scale; two anterior pairs in contact with lower labials, third pair separated by a single flat scale. Pregulars (Ruibal, 1952) flat, of various sizes and shapes. Two, larger than the others, can be considered a fourth pair of chin shields. A granular gular crease. Gulars in 9-10 transverse rows. Collar weak. Ventrals imbricate, keeled, in longitudinal and 21-24 (mean 22.7) transverse rows. Dorsals imbricate, leaf-shaped, keeled, in diagonal rows and 31-35 (mean 32.6) transverse rows. Six preanal scales; two on midline, of which one is anterior and the other is marginal; these flanked by two large marginal scales; occasionally, only a single elongate, median preanal scale rather than two. Adult males with 2-2 preanal pores and 10-14 (mean 11.7) total femoral pores. Adult females with 2 faint preanal pores but no femoral pores. Color and markings much like those of L. percarinatum. Body very light brown, with two thin, irregular, dark dorsolateral bands on neck, shoulders, and back, bordered above by distinct white stripes in young individuals and adult males. Dark bands and white stripes both fade in mature females. L. guianense lacks the distinct white stripe on the tail found in L. percarinatum, although some specimens have faint traces of it. Labials and chin shields with light brown spots; ventral surface white, side of neck with several faint white spots; only a faint white spot on frenoocular. 16 POSTILLA 154 REMARKS. The holotype (UMMZ 46770) and paratype (UMMZ 46768) were collected by Alexander G. Ruthven at Dunoon, Demerara, Guyana on 27 July 1914. Ruibal (1952: fig. 1) emphasized the arrangement of the preanal scales as diagnostic of Leposoma guianense. Only five of the specimens examined show this arrangement (UMMZ 46768, 46770, BMNH 1930.10.10. 168, 1968.877, MNHN 99.70). In FMNH 30935, the middle preanal is elongate but not as long as the paramedian preanals; there is only one median scale, however. In general, the preanal scales in L. guianense are rela- tively longer than the preanal scales of L. percarinatum. A long median preanal scale occurs in some specimens of L. parietale (USNM 166351, UKMNH 121815) although it has not been observed in L. percarinatum. SPECIMENS EXAMINED. Brazil: Amapa: Ponta dos Indios (0-100 m) BMNH 1938.3.2.2; upper Rio Calgoene (Rio Carsevenne) (100-200 m) MNHN A L. southi WL soonna MAP 2. Northwestern South America showing localities for specimens of Leposoma parietale, L. ioanna, new species, and Colombian records for L. southi. Certain localities discussed in the text are also indicated. LIZARDS OF THE GENUS LEPOSOMA i | 98.182; upper Rio Lunier, Tumuc-Humac (Tumucumaque; 100-200 m) MNHN 99.68, 99.69, 99.70 (2 specimens); Serra do Navio (100 m) UKMNH 97868. French Guiana: Inini: Camopi (100-200 m) MNHN 02.261-264. Guyana: Demerara: Dunoon (0-100 m) UMMZ 46768, 46770 (paratype and holotype of Leposoma guianense); Essequibo: Morabali Creek (0-100 m) BMNH 1930.10.10.168; Winiperu (0-100 m) BMNH 1968.877. Surinam: Nickerie: Frederick Willem IV Falls (112 m) FMNH 30935; Saramacca: near Coppename River, Camp | (0-100 m) BMNH 1946.4.3.99. DISCUSSION. Ruibal (1952) considered the possibility that L. percarinatum and L. parietale (a species from the Amazonian slopes of the Andes in Peru, Colombia, and Ecuador; Map 2) are conspecific. If L. percarinatum is a unisexual species, this seems an unlikely possibility. Unisexual species usually arise as a result of hybridization (Uzzell, 1970a). What two species may have hybridized to produce L. percarinatum? Leposoma guianense and L. parietale seem likely candidates. In Figures 8 and 9 and in Table 1, certain features of L. parietale, L. guianense, and L. percarinatum are compared. In some of these (number of dorsal scale rows, number of scales around midbody, size of third and fourth pairs of chin shields) L. parietale and L. guianense are very similar. In contrast, the relative size of the interparietal in L. percarinatum and L. parietale is very similar. The og Leposoma parietale @ Leposoma joanna + meanfor ZL. guianense Leposoma guianense MEAN PARIETAL LENGTH(mm) 13 its} 1.7 1.9 2.1 23 2.5 CES eg 3.1 INTERPARIETAL LENGTH (mm) FIG. 8. Mean parietal length as a function of interparietal length in Leposoma parietale. The confidence ellipse includes 95% of measurements of L. guianense. 18 POSTILLA 154 ©9 Leposoma parietale ® Leposoma ioanna + mean for L. guianense Leposoma guianense 45 INTERPARIETAL AREA (mm?) Gin as ol (oe) w {e) fw) e) 20 IS) 20) 22> 24526) 28" S50 SZ) Ss 4 SCO OmmO SNOUT-VENT LENGTH (mm) FIG. 9. Approximate interparietal area as a function of body length in Leposoma parietale. The oval includes all measurements of L. guianense. size of the scale behind the longest supralabial in L. parietale is intermediate between the conditions seen in L. percarinatum and L. guianense. These data do not prove that L. percarinatum, if truly a unisexual species, arose as a result of hybridization of L. guianense and L. parietale, but the data suggest this to us. The mean for the specimens of L. percarinatum is intermediate between the means for L. guianense and L. parietale for the second canonical variate, which reflects primarily features of the interparietal scale (Fig. 8). The mean is closer to that for L. parietale, even though L. percarinatum is sympatric with L. guianense. In the first canonical variate, L. percarinatum is unlike either bisexual species. Other combinations (for instance, the possibility that L. rugiceps is a parental species) seem less likely on both geographic and morphological grounds. We would argue that a clear demonstration (through karyotypes or through electrophoretic mark- ers) that L. percarinatum represents a hybrid of L. guianense and L. parietale would be a compelling argument that L. guianense and L. parietale are not conspecific. That hybrids between the two are so imbalanced genetically that one sex is repressed and the meiotic mechanism is modified, as it appears to be in all unisexual species (Uzzell, 1970a), would be, we feel, strong LIZARDS OF THE GENUS LEPOSOMA 19 evidence that L. parietale and L. guianense are so distinct genetically that antihybridization mechanisms (Remington, 1968) would be selected for if the two were in contact. Morphological differences, whether on a gross or a molecular level, cannot usually support this judgment for allopatric populations. The distribution of unisexual taxa provides additional evidence about the localization of areas of extensive recent hybridization to supplement the summary offered by Remington (1968). The following description of L. parietale is provided for comparison with the descriptions of L. guianense and L. percarinatum. Leposoma parietale (Cope) Mionyx parietalis Cope, 1885, Proc. Amer. Philos. Soc. 23: 96. DESCRIPTION. Head broad; dorsal head scales longitudinally striate. Fron- tonasal single, wider than long, separated from anterior supraoculars. Nasal divided, slightly elongate posteriorly. Loreal divided; upper segment higher than long; lower segment large. Prefrontals shorter than frontonasal. Frontal long and hexagonal. Frontoparietals elongated back along posterior supra- oculars, joined by a moderately long suture. Interparietal large, hexagonal, generally long and narrow. Parietals smaller and shorter than interparietal; anteriormost part of each parietal usually projecting forward to about the same level as anteriormost part of interparietal. Supraoculars 4; superciliaries 4; suboculars 4-6. Temporals small, keeled. Upper labials 6-7, the sixth the longest, the seventh small, but larger than adjacent temporals. Lower labials 5—6, the third the longest. A mental followed by one undivided and four paired chin shields; two anterior pairs of chin shields in contact medially, posterior two pairs separated by several small scales; two anterior pairs in contact with lower labials. Gulars in 9-10 transverse rows, the first row separated from chin shields and pregulars (Ruibal, 1952) by a row of granular scales. Collar indistinct. Ventrals imbricate, keeled, in longitudinal and 20—24 (mean 22.3) transverse rows between collar and preanal scales. Dorsals imbricate, leaf-shaped, in diagonal and 29-36 (mean 33.1) transverse rows between interparietal and posterior margins of hind limbs. Caudals imbricate, in longitudinal and transverse rows. Six preanal scales; usually two on midline, of which one is anterior and the other marginal, these flanked by two large marginal scales, which in turn are flanked by two smaller scales, also marginal. Males with 2-2 preanal and 11-17 (mean 14.4) total femoral pores. Many (56%) females with 2 preanal pores; no females with femoral pores. Limbs slender, digits also. First digit smallest, fourth largest, second and fifth shorter than third. On pes the fifth is longer than second. Dorsum usually brown, rather dark in some individuals; sides of neck and 20 POSTILLA 154 body heavily peppered with dark concentrations of pigment. Ventral surface cream. Side of neck with several white spots; labials with dark spots on their centers; lower part of loreal with white spot. Juvenile specimens and many adults with a series of white spots in two dorsolateral rows beginning on neck just behind head and continuing down over shoulders to base of tail; these spots may fuse into dorsolateral streaks. The streaks are usually most marked anteriorly, but some specimens have only faint traces of them. VARIATION. These is some variation in coloration. The main differences are in intensity, rather than distribution. The dark color of certain areas is the result of small dots of pigment set in the white epidermis. Variations in intensity are related to the density of dots of pigment underneath the scales. The brown color is in certain of the scales, rather than the epidermis. White areas, such as the ventral surfaces and dorsolateral stripes, are covered by whitish, translucent scales. Ranges of counts of dorsal and ventral scales and scales around midbody region in L. parietale are given in Table 1. Certain individuals have a com- bination of high dorsal and high ventral scale counts. None of these in- dividuals (MCZ 86644; USNM 166343, 166353; UKMNH 109771, 112208, 121819, 121822, 121825) shows any special affinities with L. percarinatum, however. Two specimens (USNM 166351; UKMNH 121815) have the distinctive five-scale anal shield described by Ruibal (1952) for L. guianense and thought by him to be characteristic of that species. Except for the number and arrangement of their preanal scales, both USNM 166351 and UKMNH 121815 are very similar to the other specimens referred to L. parietale. Among the 106 specimens examined there were 62 males and 39 females. REMARKS. The holotype of L. parietale was collected at Pebas, Loreto, Peru, by John Hauxwell sometime between 1870 and 1885. Pebas remains the easternmost locality for the species. The holotype was probably in the Academy of Natural Sciences of Philadelphia. If so, it has apparently been lost (E. V. Malnate, personal communication). The specimens examined agree with Cope’s description except that the holotype had smooth head scales, a straight rudimentary claw on the first digit of the manus, and a fifth digit on the pes that was smaller than the second. As noted by Ruibal (1952) these differences may be based in part on the poor condition of the holotype reported by Cope. SPECIMENS EXAMINED. Colombia: Caqueta: Morelia (500 m) ANSP 25507-8; Villa Maria, 45 km S of Florencia (200-500 m) MCZ 61156; Putumayo: 10 km S of Mocoa (7-800 m) AMNH 106635—36; 7 km SE of Mocoa (700 m) AMNH_ 106637. Ecuador: Morona-—Santiago: Lagarto Cocha (200-500 m) USNM 166355-56; Taisha (510 m) USNM 166357; Napo: Braga Cocha MCZ 93416-17; SE foot of Cordillera de Galeras, upper Rio LIZARDS OF THE GENUS LEPOSOMA Z1 Napo (600 m) USNM 166347; Limon Cocha (200-500 m) UKMNH 98601- 03; Loreto (500-1000 m) USNM 166344—46; Puerto Libre, Rio Aguarico (570 m) UKMNH 121826, 121852-54; 121784-808; Puerto Ore, Rio Aguarico (420 m) UKMNH 121810; Rio Llushin, N of Arapicos (500— 1000 m) USNM 166348; Rio Pucuno (500-1000 m) USNM 166343; San Francisco, Rio Napo (200-500 m) UMMZ 84740-41; San José Viejo de Sumaco (+ 1100 m) USNM 166349-S0; Santa Cecilia (340 m) UKMNH 105368-69, 107035—36, 109767—72; 112205-16, 121811-25, 121827-31; near Tena (500 m) UMMZ 84739; 15 km ENE of Umbaqui (530 m) UKMNH 121809; Pastaza: Montalvo (314 m) USNM 166351-53; Sarayacu (400 m) MCZ 38442; USNM 166354. Peru: Loreto: Iquitos (117 m) AMNH 56260; FMNH 45477; near Iquitos, Rio Itaya (100-200 m) AMNH 56261-62. III. Leposoma ioanna, A NEW SPECIES FROM PACIFIC COASTAL COLOMBIA, AND RECORDS FOR Leposoma southi, NEW TO THE FAUNA OF COLOMBIA Eleven specimens of Leposoma have been examined from the Pacific low- lands of Colombia. Two of these resemble L. parietale of the Amazonian slopes of Peru, Ecuador, and Colombia. Although the morphological dis- tinctness of these two specimens is slight, they have probably been isolated from the Amazonian population by the Andes for sufficiently long so that they would develop antihybridization mechanisms (Remington, 1968) were contact to be renewed. I therefore name them as a distinct species. Leposoma ioanna, new species HOLOTYPE: FMNH 165798, an adult female from a lumber camp, “Carton de Colombia,” 7 km W of Rio Calima, Valle del Cauca, Colombia, on road to Buenaventura; 0-500 m above sea level; collected August 1963 by Isidoro Cabrera; original number 44. PARATYPE: FMNH 165804, an adult female from 22 km on road from Buenaventura to Rio Calima, Valle del Cauca, Colombia; 0-500 m above sea level; collected April 1962 by Isidoro Cabrera; original number 82. DIAGNOSIS: A member of the genus Leposoma characterized by a single frontonasal, ventrals arranged in longitudinal rows, and conical rather than leaf-shaped scales on the side of the neck. The first feature distinguishes L. ioanna from L. southi, L. scincoides and L. annectans; the second distin- guishes it from L. scincoides and L. annectans, and the third distinguishes it from L. rugiceps (Fig. 10). Leposoma ioanna is most closely related to L. percarinatum, L. guianense, and especially L. parietale. It differs from L. percarinatum in having an 22 POSTILLA 154 FIG. 10. Ventral view of the heads of specimens of Leposoma parietale and Leposoma ioanna, new species (10). Above, UKMNH 121825, adult female L. parietale; below, FMNH 165798, adult female, holotype of L. ioanna. In L. ioanna, the third pair of chin shields is much reduced, the fourth pair is absent; the third pair is relatively large and a fourth pair is present in L. parietale. LIZARDS OF THE GENUS LEPOSOMA 25 FIG. 11. Dorsal and oblique ventral views of the head of Leposoma ioanna, new species (X10). The oblique view shows the two small scales separating the much reduced third paired chin shield from the infralabials. 24 POSTILLA 154 enlarged scale behind the longest supralabial; in having fewer (28-30) transverse dorsal scale rows (36-39 in L. percarinatum), and the third pair of chin shields much reduced and separated from the infralabials by two small scales (Figs. 10, 11). It differs from L. guianense in having the third pair of chin shields much reduced and separated from the labials by 2 small scales, a relatively shorter interparietal compared to the parietal length, and fewer transverse dorsal scale rows (31-35, mean 32.6 in L. guianense). It is most like L. parietale, from which it differs in having the third pair of chin shields much reduced and separated from the infralabials by two scales, and a somewhat reduced dark band (approximately two scales wide) along the sides of the body (4—S scales wide in L. parietale). It may also average fewer transverse dorsal scale rows (29-36, mean 33.5 in L. parietale) and a larger maximum size (largest female of L. parietale measured was 38 mm snout-to-vent; holotype and paratype of L. ioanna are 41 and 39 mm snout- to-vent, respectively). DESCRIPTION OF HOLOTYPE: (Figs. 10, 11). Rostral in contact with first supralabial, nasal, and single frontonasal. Frontonasal wider than long, in contact with rostral, nasal, loreal (narrowly) and paired prefrontals. Frontal approximately twice as long as broad, followed by paired frontoparietals. Interparietal longer than broad. Interparietal length 1.25 times parietal length. Supraoculars 4—4, the third largest, the first and fourth smaller. Superciliary series complete. Nasal divided into two parts behind the nostril. Loreal divided, the upper part small, in contact with lower part, nasal, frontonasal, prefrontal, large first superciliary, and a minute frenoocular; lower part in contact with second and third supralabials. A complete series of very small infraorbital scales. Supralabials 6-6, the last longest and highest, almost completely behind the orbit. Scale behind 6th supralabial along lip line noticeably larger than adjacent temporal scales. Temporal region covered with small scales. Scales of dorsal surface of head with longitudinal striations except for rostral and frontonasal. Scales of side of head smooth anterior to orbit, with longitudinal ridges posterior to orbit. Infralabials 5—S. A mental; a single unpaired chin shield, followed by two large paired chin shields that contact each other across the midline and that contact the infralabials. Third pair of chin shields much reduced, separated from each other by small scales and from infralabials by two small scales (Figs. 10, 11). A gular fold with granules; pregulars (Ruibal, 1952) weakly keeled, obtusely angled. Gulars strongly keeled, long pointed, in diagonal and 7 longitudinal rows, including the weakly developed collar. Dorsal scales immediately behind parietals and interparietal irregular; narrow, elongate, and strongly keeled on neck; becoming leaf-shaped and strongly keeled on dorsal surface of body; imbricate, in 28 irregular transverse rows between interparietal and posterior margin of hind limbs. Scales on side of neck conical; in axilla and groin, granular. Along sides of body, scales like dorsals but smaller and more strongly keeled. Lateral scales LIZARDS OF THE GENUS LEPOSOMA 22 grading into ventral scales, which are keeled and mucronate. Twenty-one scales around the midbody region. Between the arms, the imbricate ventral scales are in diagonal and transverse rows; behind this, in longitudinal and transverse rows. There are 20 transverse rows between the collar and the two rows of preanal scales. Five weakly keeled scales along the posterior margin of the vent, the paramedian larger: anterior to these, another row with a large median scale and two smaller, more anterior paramedian scales. Scales of tail in complete rings, those on dorsal and lateral surfaces more strongly keeled. Preanal pores represented by one minute chink on each side. Scales of limbs keeled, overlapping, smallest on ventral surface of upper forelimb and on posterior surface of thigh. Scales at base of palm slightly enlarged; other palmer and plantar scales small, conical. Subdigital lamellae apparently doubled. Coloration. The dorsal surface of the head is heavily mottled with dark brown. Anterior dorsal surface of snout lighter. Side of head dark but with light areas at sutures between supralabials. Infralabials dark spotted, but chin shields only lightly sprinkled with dark pigment, so that midventral area of head is light. Ventral surface of body generally light. Underside of tail with some dark pigment posteriorly. Laterally, an irregular dark brown dorsolateral line about two scales wide extending onto the tail. Above this a much lighter tan dorsal area, sprinkled middorsally with dark brown pig- ment. A dorsolateral light line above sacrum. No evidence of a light line along side of tail from hind limb insertion. Body length 41 mm; total length (tail tip regenerated) 95 mm. Variation. The paratype is similar in most respect, although with generally darker coloration. The paratype has 20 transverse rows of ventral scales between the collar and the preanal scales, 22 scales around the midbody region, and 30 transverse rows of dorsal scales. It is 39 mm snout-to-vent. The tail is missing. The area of interparietal, estimated as mean of anterior and posterior widths times length, is 4.1 mm for the holotype, 3.9 mm for the paratype. Several features of the holotype and paratype of L. ioanna were used in a canonical analysis along with specimens of closely related species (Fig. 7). This analysis confirms the distinctness of L. ioanna as far as it is possible on this sample. The name ioanna is from the Greek wavva; it is feminine and used in apposition to the generic name. Leposoma southi Ruthven and Gaige Alopoglossus copii, Boulenger, 1913, Proc. Zool. Soc. London, p. 1033. Two specimens (BMNH 1913.11.12.35—36) collected by H. G. F. Spurrell at Pefia Lisa (120 m elevation), near Condoto, Choc6, Colombia, were re- 26 POSTILLA 154 ported by Boulenger (1913) as Alopoglossus copii. Each has the surface of the tongue covered with diamond-shaped papillae, rather than with the oblique folds that characterize Alopoglossus. Boulenger’s identification em- phasizes the morphological similarity of Leposoma and Alopoglossus (Ruibal, 1952; Uzzell, 1969b). In addition to the two British Museum specimens, several other specimens not distinguishable from Leposoma southi have been examined (Table 1). They resemble L. southi and differ from L. rugiceps in having a divided fronto- parietal and in having conical rather than leaf-shaped, imbricate scales on the side of the neck (Fig. 12). There is no light line along the side of the tail beginning at the hind limb insertions; such a light line is present in L. rugiceps. In other respects, the specimens fall in or near the limits of variation reported by Ruibal (1952). There are 19-21 (mean 20.1) transverse rows of ventral scales between the collar and preanal pores, 27—29 (mean 28.2) dorsals between the parietal and the posterior margin of the hind limbs, and 21—23 (mean 21.6) scales around the midbody region. The third pair of postmentals are separated from the labials by two small scales, and are not in contact on the midline. The three males have 2—2 preanal pores FIG. 12. Side of the neck of Leposoma rugiceps (MLS 716) showing imbricate scales resembling dorsals (X11). LIZARDS OF THE GENUS LEPOSOMA pa | and 10-11 (mean 10.3) total femoral pores. The six females have only a single preanal pore on each side. The longest male is 40 mm snout-to-vent, the longest female, 37 mm. BMNH 1913.11.12.35 has 7 rather than 6 supra- labials on the left side. The scale posterior to the long, last supralabial is relatively large, as in L. guianense (Fig. 5). Ruibal (1952) considered the possibility that L. southi and L. rugiceps, which appeared to have allopatric ranges, are conspecific. He found no evi- dence of hybridization in the areas where the ranges of the two species ap- peared to approach each other. L. southi previously was known only from southern Costa Rica and in Panama east to Barro Colorado in the Canal Zone. Specimens have since been collected in eastern Panama (William E. Duellman, personal communication). L. rugiceps is known from the Canal Zone (but not Barro Colorado) east to the Santa Marta region of Colombia, and south in the valley of the Rio Magdalena (Ruibal, 1952). Taylor (1955, 1956) recognized two subspecies of L. southi in Costa Rica. Their relations to each other, to Panamanian populations, and to the Colom- bian specimens are not clear. SPECIMENS EXAMINED. Colombia: Choc6: Pena Lisa, Condoto (300 m) BMNH 1913.11.12.35—36; 2 km above Playa de Oro, upper Rio San Juan (210 m) AMNH 107264—65; Quebrado Docord6, ca. 10 km above junction with Rio San Juan (100 m) AMNH 107263; Quebrado Vicordo, ca. 5 km above Noanama, Rio San Juan (80-110 m) AMNH 107259-62. DISCUSSION. Discovery of populations of two species of Leposoma on the Pacific coast of Colombia reveals how much has yet to be learned about the herpetofauna of South America. The discovery of L. ioanna and the inter- pretation of L. percarinatum as a possible hybrid between L. guianense and L. parietale both suggest considerable shifting of the ranges of these lizards during their history. Leposoma ioanna clearly shares a common ancestor with L. parietale. On the other hand, these two species are presently separated by the Andes, and the lowland area at the north end of the Andean ranges is occupied by L. rugiceps. It does not seem possible that they could recently have been con- nected by continuous populations. The events that account for the present separation, which is surely real, are not known. If L. percarinatum is correctly interpreted as a species resulting from hy- bridization of L. parietale and L. guianense, then those two species, which appear very similar, must have been in contact at some time in the past. The most plausible history for these forms is disjunction of an ancestral popula- tion, probably across the lowlands between the Guianan Highlands and the Amazonian Andean slopes, followed by speciation. After some interval, the two daughter species again came into contact and, as a result of hybridization, produced the species L. percarinatum. At present, however, L. parietale and L. guianense seem no longer to be in contact. 28 POSTILLA 154 Although these events are speculative, it would be of great interest to es- tablish them more firmly, and then to establish probable dates for the various events. The persistence of many unisexual species would seem to be prolonged by a fixed heterozygosity that to some extent vitiates the absence of recom- bination, but unless sexuality can be restored, such species appear to be doomed in an evolutionary sense. IV. A KEY TO THE SPECIES OF Leposoma The following key is modified from Ruibal’s 1952 key in light of the speci- mens reported here: 1. Frontonasal longitudinally divided ©... 3). Z Frontomasal single: oo eccccocg ceccsde ieee ate ode snes eee + 2. Pregulars flat and quadrangular; ventrals in longitudinal rows; range: Costa Rica, Panama, and Pacific Colombia ........................5 L. southi Pregulars convex, posteriorly pointed; ventrals not in longitudinal TOWS © see rissa teense nvdnnn cia tiealsieres aa sioner ined teetynd gaelic eee nee ee 3 3. Interparietal neither longer nor broader than the parietals; third post- mental in contact with the lower labials; range: Brazil ............................ patra etches ee AO mentee aR et re A ou ee ea L. scincoides Interparietal longer and broader than parietals; third postmental sep- arated from the lower labials by a small scale; range: Baia, Brazil ............ Beg Wa eautet ead Mace cce raat oh Sen hi ten to eGo oe RO ae eee L. annectans 4. Scales on side of neck keeled, imbricate, shaped like dorsals; 27-31 scales from parietals to posterior margin of hind limbs .......... L. rugiceps Scales on side of neck not shaped like dorsals; 28-39 scales from parietals to’ posterior margin of hind limbs ~....7..05.5) =e eee 5) 5. An irregular dorsolateral dark stripe 1 or 2 scales wide, extending onto tail where it is bordered below by a white stripe originating at insertion of hind limb; 36-39 scales from parietals to posterior margin of hind limbs; fourth pair of chin shields very small; no males known; Guianas, adjacentsVenezuclasand Brazilis. ee eee L. percarinatum Dorsal scales 28-36; fourth pair of chin shields very small or moderately Lar Bey fe eet lve Renae: Saaceisccee ena eet eee 6 6. Dark lateral stripe 4-5 scales wide; fourth pair of chin shields moder- ately large; Amazonian slopes of Ecuador and adjacent Colombia and Berg oe oc as et ee ie Bd nh eee eT eee L. parietale Dark lateral stripe 1-2 scales wide; fourth pair of chin shields moderate Or reduced! .i 0. 6An. ce eee tees eee ii 7. Fourth pair of chin shields moderately large; Guianas and adjacent 18) 217A | Ue eames ei oe OMRON TRE RAIS re PRR i hes Caan EN FB Wiehe ey SEE To 0 L. guianense Fourth pair of chin shields very small; Pacific Colombia ........................ | LIZARDS OF THE GENUS LEPOSOMA 29 V. THE HEMIPENIS OF SPECIES OF Leposoma Hemipenes of three species of Leposoma have been examined. The left organ was removed from the lizard, washed overnight in distilled water, and stained with alizarin red S in 1% KOH. The hemipenes were slit along the sulcus spermaticus. In Leposoma rugiceps (Fig. 13) the inverted organ has almost no median welt. The right and left halves contain well marked W-shaped flounces. At each apex of the W, there is a single large calcareous tooth. In both the right (medial) and left (lateral) halves of the organ, there are thus two striking rows of teeth; these number about 9 on each half. The distal part of the organ is marked by fleshy lobes. There are no calcareous spines in the basal part of the organ. No everted organs have been observed. ad a ot , * k * ‘ioe “a oe, ahi? : * ay é FIG. 13. Structure of the left hemipenis of Leposoma rugiceps (BMNH 1929.6.1.84). The inverted organ has been slit along the sulcus spermaticus and laid open, show- ing two rows of calcareous spinules in both lateral (left) and medial (right) pockets (32). 30 POSTILLA 154 FIG. 14. Hemipenis of Leposoma guianense (BMNH 1698.877). A, left organ slit along sulcus spermaticus and laid open, showing two rows of calcareous teeth in each pocket (X25). B, C, details of teeth (x36). | LIZARDS OF THE GENUS LEPOSOMA BS The hemipenis of L. guianense is similar in many respects to that of L. rugiceps. The median welt is poorly developed, and there are weak W-shaped flounces in both medial and lateral halves of the organ. At the two apices of each flounce, there is a single large calcareous tooth (Fig. 14.) The teeth, which number between 12 and 14, are unicuspate, although some of them have broad bases (Fig. 14B, C) and development of cusps on these teeth in some individuals is easy to imagine. The hemipenis of L. parietale (Fig. 15) also has a reduced median welt and W-shaped flounces in both medial and lateral halves. The two apices of each W bear a single large unicuspate tooth, although many of these have broad bases. There are more teeth (23—26) per row of apices than in L. rugiceps and L. guianense. In marked contrast to the hemipenes of L. rugiceps and L. guianense the flounces of the hemipenis of L. parietale contain series of denticles extending out from the apical tooth towards the median welt and towards the sulcus spermaticus. These denticles decrease in size with greater distance from the apical tooth (Fig. 15B). Denticles seem to be absent between the two apical teeth of a single W. These denticles lack the orderliness seen in the calcareous spinules in the flounces of many species of group II of the family Teiidae (cf. Uzzell, 1969a: figs. 2,3; 1970b: figs. 3-5). The hemipenes of the three species that have been examined show certain common features. The median welt is reduced, and both the medial and lateral lobes of the organ have W-shaped flounces. Large teeth on the two apices of each W are common to all three species; this is the condition reported previously (Uzzell, 1969b). The hemipenis of L. parietale shows, in addition to these two rows of teeth, numerous denticles in each flounce. In this re- spect, the hemipenis shows a marked similarity to the hemipenis of Arthro- saura kockii (Figs. 16, 17). These flounces number about 20—23. On each flounce, extending from the enlarged tooth towards the median welt or from the other enlarged tooth towards the sulcus spermaticus, there is a long row of denticles. Such denticles also occur on some of the flounces between the large teeth, but are fewer and less well developed (Fig. 17). On the basis of numerous features of scalation, Arthrosaura and Leposoma appear to be related (Uzzell, 1969b). The similarity of the hemipenial spines and flounces of L. parietale and A. kockii, especially the weak median welt, the W-shaped flounces in each half of the organ, the well developed teeth at the two apices of each W, and the numerous denticles adjacent to these strong teeth support the relationships of these two genera. ACKNOWLEDGEMENTS For use of specimens in their charge, we thank James E. Bohlke, Academy of Natural Sciences of Philadelphia, William E. Duellman, University of Kansas Museum of Natural History, Alice G. C. Grandison, British Museum (Natural History), Jean Guibé, Museum National d’Histoire Naturelle, Robert F. Inger 32 POSTILLA 154 LIZARDS OF THE GENUS LEPOSOMA 33 and Hymen Marx, Field Museum of Natural History, Hno. Nicéforo Maria, Museo de La Salle, Bogota, James A. Peters, U. S. National Museum, Charles F. Walker, University of Michigan Museum of Zoology, Ernest E. Williams, Museum of Comparative Zoology, Harvard University, and Richard G. Zweifel, American Museum of Natural History. Edmund V. Malnate, Acad- emy of Natural Sciences of Philadelphia, and Konrad Klemmer, Sencken- bergische Museum, Frankfurt-am-Main kindly answered questions about particular specimens in those collections. Charles W. Myers, American Mu- seum of Natural History, allowed us to report on his specimens of Leposoma from Colombia. Photographs were made by A. H. Coleman, and figures by Rosanne Rowen. The senior author particularly thanks Robert Mertens, Senckenbergische Museum, Alice G. C. Grandison and E. N. Arnold, British Museum (Natural History), and Jean Guibé for hospitality while visiting their particular institutions. Computer time was available through NSF grant GB 8769 to the senior author. Travel in Europe by the senior author was made possible by Yale University and a gift of Evan Commager. FIG. 15. Hemipenis of Leposoma parietale (USNM 116351). A, left organ slit along sulcus spermaticus and laid open (24). B, C, details of large teeth on apices of flounces (X33). D, detail of enlarged teeth and of denticular series extending along edges of flounces away from enlarged teeth (33). POSTILLA 154 34 LIZARDS OF THE GENUS LEPOSOMA 35 FIG. 16. Left hemipenis of Arthrosaura kockii (USNM 154269) slit along sulcus spermaticus and laid open to show lateral (left) and medial (right) pockets (x 23). POSTILLA 154 36 i ey LIZARDS OF THE GENUS LEPOSOMA a7 FIG. 17. Medial pocket of left hemipenis of Arthrosaura kockii (USNM 159269), showing details of the enlarged, uni- to tricuspid teeth and the denticular rows along the edges of the flounces (38). 38 POSTILLA 154 LITERATURE CITED Boulenger, George Albert. 1885. Catalogue of the lizards in the collection of the British Museum, II. Taylor and Francis, London. 497 p. 24 pl. 1913. On a collection of batrachians and reptiles made by Dr. H. G. F. Spurrell, F. Z. S., in the Choco, Colombia. Proc. Zool. Soc. London 1913: 1019-1038. Burt, Charles E., and May Danheim Burt. 1931. South American lizards in the collection of The American Museum of Natural History. Bull. Amer. Nat. Hist. 61 (7): 227-395. Cope, Edward Drinker. 1885. Catalogue of the species of batrachians and reptiles contained in a collection made at Pebas, upper Amazon, by John Hauxwell. Proc. Amer. Philos. Soc. 23: 94-103. Cunha, Osvaldo Rodrigues da. 1961. Lacertilios da Amaz6nia. II. Os lagartos da Amazonia Brasileira, com especial referéncia aos representados na colecao do Museu Goeldi. Bol. Mus. Paraense Emilio Goeldi, n.s., Zoology, no. 39, 189 p. Darevsky, Ilya S., and V. N. Kulikova. 1961. Natiirliche Parthenogenese in der polymorphen Gruppe der kaukasischen Felseidechse (Lacerta saxicola Evers- mann). Zool. Jahrb. Syst. 89: 119-176. Dixon, W. J., ed. 1968. BMD Biomedical Computer Programs. Univ. California. Publ. Automatic Computation, No. 2. Univ. California Press, x + 600 p. Maslin, T. Paul. 1962. All-female species of the lizard genus Cnemidophorus, Teiidae. Science 135: 212-213. Mertens, Robert. 1925. Zwei neue Eidechsen aus Venezuela. Senckenbergiana 7: 75-78. Minton, Sherman A., Jr. 1958 (1959). Observations on amphibians and reptiles of the Big Bend region of Texas. Southwestern Nat. 3: 28-54. Miiller, Lorenz. 1923. Neue oder seltene Reptilien und Batrachier der Zoologischen Sammlung des bayrischen Staates. Zool. Anz. 57 (7 & 8): 145-156. Noble, Gladwyn Kingsley. 1923. New lizards from the Tropical Research Station, British Guiana. Zoologica 3 (15): 301-305. Remington, Charles L. 1968. Suture-zones of hybrid interaction between recently joined biotas. Evol. Biol. 2: 321-428. Ruibal, Rodolfo. 1952. Revisionary studies of some South American Teiidae. Bull. Mus. Comp. Zool., Harvard Coll. 106 (11): 477-529. Taylor, Edward Harrison. 1955. Additions to the known herpetological fauna of Costa Rica with comments on other species. No. 2. Univ. Kansas Sci. Bull. 37, 1 (13): 499-575. 1956. A review of the lizards of Costa Rica. Univ. Kansas Sci. Bull. 38, 1 (1): 3-322. Tinkle, Donald W. 1959. Observations on the lizards Cnemidophorus tigris, Cnemi- dophorus tesselatus and Crotaphytus wislizeni. Southwestern Nat. 4 (4): 195— 200. Uzzell, Thomas. 1969a. A new genus and species of teiid lizard from Bolivia. Postilla (Peabody Mus. Nat. Hist., Yale Univ.) 129: 1-15. 1969b. The status of the genera Ecpleopus, Arthroseps, and Aspidolaemus (Sauria, Teiidae). Postilla (Peabody Mus. Nat. Hist., Yale Univ.) 135: 1-23. | LIZARDS OF THE GENUS LEPOSOMA 39 1970a. Meiotic mechanisms of naturally occurring unisexual vertebrates. Amer. Nat. 104: 433-445. 1970b. Teiid lizards of the genus Proctoporus from Bolivia and Peru. Pos- tilla (Peabody Mus. Nat. Hist., Yale Univ.) 142: 1-39. Zweifel, Richard G. 1965. Variation in and distribution of the unisexual lizard, Cnemidophorus tesselatus. Amer. Mus. Novitates 2235: 1-49. =! : REVIEW STYLE FORM TITLE ABSTRACT NOMENCLATURE ILLUSTRATIONS FOOTNOTES TABLES REFERENCES HOR’S COPIES PROOF COPYRIGHT INFORMATION FOR AUTHORS The Publications Committee of the Peabody Museum of Natural History reviews and approves manuscripts for publication. Papers will be published in approximately the order in which they are accepted; delays may result if manuscript or illustrations are not in proper form. To facilitate review, the original and one carbon or xerox copy of the typescript and figures should be submitted. The author should keep a copy. Authors of biological papers should follow the Style Manual for Biological Journals, Second Edition (Amer. Inst. Biol. Sci.). Authors of paleontological manuscripts may choose to follow the Sugges- tions to Authors of the Reports of the U.S. Geological Survey, Fifth Edition (U.S. Govt. Printing Office). Maximum size is 80 printed pages including illustrations (= about 100 manuscript pages including illustrations). Manuscripts must be typewritten, with wide margins, on one side of good quality 81%4 x 11” paper. Double space everything. Do not underline anything except genera and species. The editors reserve the right to adjust style and form for conformity. Should be precise and short. Title should include pertinent key words which will facilitate computerized listings. Names of new taxa are not to be given in the title. The paper must begin with an abstract. Authors must submit com- pleted BioAbstract forms; these can be obtained from the Postilla editors in advance of submission of the manuscripts. Follow the International Codes of Zoological and Botanical Nomen- clature. Must be planned for reducton to 414 x 7” (to allow for running head and two-line caption). If illustration must go sideways on page, reduction should be to 4 x 714”. All illustrations should be called “Figures” and numbered in arabic, with letters for parts within one page. It is the author’s responsibility to see that illustra- tions are properly lettered and mounted. Captions should be typed double-spaced on a separate page. Should not be used, with rare exceptions. If unavoidable, type double-spaced on a separate page. Should be numbered in arabic. Each must be typed on a separate page. Horizontal rules should be drawn lightly in pencil; vertical rules must not be used. Tables are expensive to set and correct; cost may be lowered and errors prevented if author submits tables typed with electric typewriter for photographic reproduction. The style manuals mentioned above must be followed for form and for abbreviations of periodicals. Double space. Each author receives 50 free copies of his Postilla. Additional copies may be ordered at cost by author when he returns galley proof. All copies have covers. Author receives galley proof and manuscript for checking printer’s errors, but extensive revision cannot be made on the galley proof. Corrected galley proof and manuscript must be returned to editors within seven days. Any issue of Postilla will be copyrighted by Peabody Museum of Natural History only if its author specifically requests it. Acme Bookbinding Co., Inc. 300 Summer Street Boston, Mass, 02210 3 2044 wuld iN : | ¢ be ks Tey ra si eyr- ft. poalegye, { Ce bas ay PAINMS SEARLS Age Oe bras. a es ry SMS WL at WO: 4 LOL FE a ‘ Re IML OR UN CCU ee Cee ie OA a LUA A Las te 44. “4 She Mtr es gars coay 19 Pom 1 4 djtoke Udst An Ses 2 Wa Ay *y Ried Wi ‘ ‘ Ad Rack etitate arb ie ate a ag ys Aer POG Ht eran ar ae fi CAV ews a RA a oy (tey