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MDHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League 31 MARCH 1992 VOLUME 28 NUMBER 1 BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY Volume 28 Number 1 March 1992 CONTENTS SIZE AND DISTRIBUTIONAL RECORDS FOR TWO SPECIES OF AGALYCHNIS (AMPHIBIA: ANURA) IN MEXICO Hobart M. Smith and David Chiszer . 1 THE NOMENCLATURAL STATUS OF UMNOPHIS GUNTHER, 1865 (REPTILIA: SERPENTES) Hobart M. Smith, George W. Byers, Kenneth L. Williams and Van Wallach . 4 DISTRIBUTIONAL AND TAXONOMIC NOTES ON SOME LIZARDS OF THE GENUS SCELOPORUS FROM MEXICO Hobart M. Smith . 8 THE NOMENCLATURAL STATUS OF CANTORIA GIRARD, 1858 (REPTILIA: SERPENTES) Hobart M. Smith, Reeve M. Bailey, Kenneth L. Williams and Van Wallach . 12 A SECOND LOCALITY FOR GEOPHIS SALLEI (REPTILIA: SERPENTES) Hobart M. Smith and David Chiszar . 16 A MEXICAN GENUS OF TROPID OPHEINE SNAKES Hobart M. Smith and David Chiszar . 19 The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc . 2643 North Charles Street Baltimore, Maryland 21218 BULLETIN OF THE Volume 28 Number 1 March 1992 The Maryland Herpetological Society Department of Herpetology, Natural History Society of Maryland, Inc. Executive Editor . Herbert S. Harris, Jr. Steering Committee Frank Groves Jeny D. Hardy, Jr. Herbert S. Harris, Jr. Library of Congress Catalog Card Number: 76-93458 Membership Rates Membership in the Maryland Herpetological Society is $16.00 per year. Foreign $20. 00/year. Make all checks payable to the Natural Histoiy Society of Maryland, Inc. Meetings The third Wednesday of each month, 8:15 p.m. at the Natural History Society of Maryland (except May — August, third Saturday of each month, 8:00 a.m.). The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 o.m. Volume 28 Number 1 March 1992 SIZE AND DISTRIBUTIONAL RECORDS FOR TWO SPECIES OF AGALYCHNIS (AMPHIBIA: ANURA) IN MEXICO Hobart M. Smith and David Chiszer Abstract Agalychnis moreleti is recorded at a maximum s-v length of 89 mm. A. callidryas at Venustiana Carranza and Mt. Ovando, Chia¬ pas, is a first record for the Central Depression of the state. On the occasion of a recent visit by Dr. John D. Lynch to the University of Colorado Museum (UCM), a number of previously unidentified amphibians from Mexico was examined, revealing two noteworthy series of Agalychnis moreleti (Dumeril) and A. callidryas (Cope), here reported. Agalychnis moreleti Two large females (UCM 39766, 52484) were taken by Thomas MacDougall in the vicinity of Vista Hermosa, municipality of Comaltepec, district oflxtlan, Oaxaca, Mexico, July-Oct., 1968, and Dec., 1970, respectively. The species has already been reported from the same locality (Duellman, 1970: 696), but both of the present specimens exceed the maximum s-v length he recorded (Duellman, 1970: 1 13) for the species, at 82.9 mm in females. The largest, UCM 39766, measures 89 mm s-v, the other 84 mm. Only A. spurrelli of Agalychnis is known to reach a greater s- v length, at 92.8 mm in females (Duellman, 1970: 124). Agalychnis callidryas. Although several records for this species were reported by Duellman (1970: 965) very near the border of Chiapas, in the state of Tabasco, none were then available for the former state. Subse¬ quently, Altig (1979: 61) reported the species from Pichucalco; Ramirez (1982: 167) from Laguna Belgica, municipality of Ocozocuautla: and Gongora (1987:29) from Lacan j a-Chansayab , municipality of Ocosingo, all in the state of Chiapas. Johnson (1989: 4, 61) considered the species as an inhabitant of the Gulf Coastal Plain and Northern Highlands physiographic provinces. All of the records for Chiapas so far available fall within those provinces. None is available for the Central Depression Physiographic Province (Johnson, 1990). Bulletin of the Maryland Herpetological Society Page 1 Volume 28 Number 1 March 1992 It is therefore of considerable interest that two specimens (UCM 39601-2) of A. callidryas were taken by Bizi Matuda of La Esperanza, Chiapas (a coffee finca where he lived and studied botany for several decades), in the Central Depression and Sierra Madre physiographic prov¬ inces. One (39601) is from Venustiano Carranza, taken in May, 1941; this locality is at about the middle of the central Depression, on its northern edge. The other specimen (39602) was taken in June, 1940, on Mt. Ovando, about 10 km NE of Acapetahua in the Sierra Madre. Both specimens are unfortunately totally discolored and extremely dehydrated. Faint vestiges of the lateral vertical lines characteristic of the species are evident in no. 39601, but are not visible in the other. Both have the typical digital webbing (Duellman, 1970: 91). scoop-shaped snout and prevomerine teeth (Duellman, 1970: 97) of the species. Each has numerous white spots scattered over the body, limbs and even on the tongue and floor of the mouth; they appear to be diseased tissues. Occurrence of the species in the Central Depression of Chiapas is not particularly open to question, since relatively low altitudes occur there in the upper drainages of the Rio Mescalapa system, specifically Rio Chiapa. Occurrence on Mt. Ovando is definitely questionable, however. The species does not occur at high altitudes (up to 960 m fide Duellman, 1970: 112) nor on Pacific slopes north of Costa Rica (ibid.). It is possible that the “Mt. Ovando” specimen came from Atlantic slopes of the Sierra Madre somewhere on the south side of the Central Depression, northeast of Mt. Ovando, but it is very unlikely that it came from the Mountain itself, which is exclusively of Pacific drainage. We interpret both specimens as documenting the presences of A. callidryas only well within the Central Depression of Chiapas. Acknowledgments We are much indebted to Dr. John D. Lynch for confirmation of the identity of the specimens here reported; to Dr. Shi Kuei-Wu for the privilege of studying the material; to Dr. William B. Lewis for facilities enabling our studies; and to Dr. Oscar Flores Villela for help with the literature. Literature Cited Altig, Ronald. 1979. Toads are nice people. Columbia, Missouri, Gates House/ Manco. 115 pp., 10 figs, (col.), ill. Page 2 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 Duellinan, William E. 1970. The hylid frogs of Middle America. Mon. Mus. Nat. Hist. Univ. Kansas, 1: 1-753, figs. 1-324, pis. 1-72 (col.). Gongora-Arones, Eleuterio. 1 987. Etnozoologia Lacandona: la herpetofauna de Lacanja-Chan- sayab. Inst. Nac. Inv. Rec. Biot., Cuad. Divulg. , (31): 1-31. Johnson, Jerry D. 1 989. A biogeographic analysis of the herpetofauna of northwest¬ ern Nuclear Central America. Milwaukee Publ. Mus. Contr. Biol. Geol., (76): i-ii, 1-66, figs. 1-29. 1990, Biogeographic aspects of the herpetofauna of the Central Depression of Chiapas, Mexico, with comments on sur¬ rounding areas. SWem Nat., 35(3): 268-278, figs. 1-3. Ramirez-Bautista, Aurelio, Oscar Alberto Flores Villela and Gustavo Casas- Andreu. 1982. New herpetological state records for Mexico. Bull. Maryland Herp. Soc., 18(3): 167-169. Department of Environmental Population and Organismic Biology, University of Colorado, Boulder, Colorado 80309-0334 (HMS); and Department of Psychology, ibid., 80309-0345 (DC). Received: 29 May 1991 Accepted: 6 June 1991 Bulletin of the Maryland Herpetological Society Page 3 Volume 28 Number 1 March 1992 THE NOMENCLATURAl STATUS OF LIMNOPHIS GUNTHER, 1865 (REPT1LIA: SERPENIES) Hobart M. Smith, George W. Byers, Kenneth L. Williams and Van Wallach Abstract Umnophis Blanchard, 1 845, is an “incorrect subsequent spelling” of Limnobia Meigen (1818). So interpreted, it has no nomenclatural status and does not invalidate the valid name Limnophis Gunther, 1865. As pointed out by Williams and Wallach (1989: 86), the currently accepted name Limnophis Gunther (1865), for a genus of central African snakes (occurring in Angola, Zaire, Botswana, Zambia and Zimbabwe), is antedated by Limnophis Blanchard (1845) applied to a genus of crane flies, family TipuHdae, for which Meigen (1818) proposed the name Limnobia. From the latter genus Macquart (1834) separated a group that he named Limnophila. However, it is clear from the original proposal that Blanchard’s (1845: vol. 2, 455) Limnophis is a lapsus for Limnobia , because the name was attributed to Meigen, who never used that name but did propose Limnobia. That Blanchard meant that name is evident from his citation Limnophila (misspelled Limnephilal ) Macquart as a synonym, inasmuch as Macquart (1834) erected that genus in partitioning Meigen’s Limnobia. Although it seems clear that Blanchard’s Limnophis is a lapsus for the intended name Limnobia Meigen, his concept of that genus was that of Macqu art’s Limnophila , as indicated by his statement that usually 16 antennal segments occur, whereas what Meigen named Limnobia (now Limonia Meigen, 1803) typically has 14 antennal segments. That misiden- tification does not alter the fact that Blanchard obviously intended to write Limnobia instead of Limnophis. The name Limnophis Blanchard has never been used as a valid name. Nomenclators (Sherbom, Schulze et al., Neave) uniformly regard the name as a substitute for (“emendation” of) Limnophila Macquart. Dipterists, Page 4 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 however, appear to have ignored the name or recognized it as a meaningless misspelling (e.g., Kertesz, 1 902) . Were the emendation interpretation upheld, Linmophis Blanchard, 1845, would have to be accepted as a nomenclaturally available name, and Linmophis Gunther would thus be a junior homonym that “must not be used as a valid name” (Art. 52a, 1985 Code), at least without resort to the plenary powers of the International Commission on Zoological Nomenclature. On the contrary, we here hold it clearly evident, for the reasons previously stated, that Blanchard’s name Linmophis was “an incorrect subsequent spelling” (Art. 33c) for Limnobia Meigen (not Linmophila Mac- quart), and that it “is not an available name and therefore does not enter into homonymy” (Art. 33c). Accordingly, Linmophis Gunther, 1865, is not antedated by any senior homonym that is available (oiz., Linmophis Blanchard, 1845), and no action is needed by the International Commission on Zoological Nomencla¬ ture to affirm the nomenclatural validity of Linmophis Gunther, so far as known at present. Also as noted by Williams and Wallach (1989: 86), the generic name Linmophiswas proposed as new by Marsh ( 1 87 1 ) for a fossil reptile. The same author, discovering that his name was preoccupied, later proposed the substitute name Lestophis (Marsh, 1885). Schmidt (1927), unaware of Marsh’s substitute name, proposed Paleoboa as a replacement for Linmo¬ phis Marsh. The valid name Lestophis remains a nomen dubiimu however, although it is referable to the Rhineuridae, a family of the reptilian order (or suborder) Amphisbaenia (Estes, 1983: 202), not to the Serpentes, as long thought. Acknowledgments We are greatly indebted to Dr. Jon K. Gelhaus of the Academy of Natural Sciences of Philadelphia for consulting and duplicating for us the pertinent parts of works not otherwise available to us. Literature Cited Blanchard, Emile. 1845. Histoire des insectes, traitant de leurs moeurs et de meta¬ morphoses en general, et comprenant une nouvelle classi¬ fication fondee sur leurs rapports naturels. Paris, Didot. 2 vols., 398, 544 pp., 20 pis. Bulletin of the Maryland Herpetological Society Page 5 Volume 28 Number 1 March 1992 Estes, Richard. 1983. Handbuch der palaoherpetologie. Teil 10A. Sauria ter- restria, Amphisbaenia. Stuttgart, Fischer, xxii, 249 pp., figs. 1- 69. Gunther, Albert C. L. G. 1 865 . Fourth account of new species of snakes in the collection of the British Museum. Ann. Mag. Nat. Hist., (3)15: 89-98, pis. 2- 3. Kertesz, Koloman. 1902. Catalogus dipterorum hucusque descriptorum. Vol. 2. Budapest, Hungarian Natl. Mus. 359 pp. Macquart, Jean. 1834. Histoire naturelle des insectes. Dipteres. Vol. 1. Paris, Roret. 578 pp., 12 pis. Marsh, Othniel C. 1871. Description of some new fossil serpents, from the Tertiary deposits of Wyoming. Am. J. Sci., (3) 1: 322-327. 1885. Names of extinct reptiles. Am. J. Sci., (3)29: 169. Meigen, Johann Wilhelm. 1803. Versuch einer neuen Gattungseintheilung der europais- chen zweiflugeligen Insekten. Mag. f. Insektenkunde, 2: 259-281. 1818. Systematische Beschreibung der bekannten europaischen zweiflugeligen Insekten. Vol. 1. Aachen, Schulz. 33 pp., 1 1 pis. Schmidt, Karl P. 1927. New reptilian generic names. Copeia, (163): 58-59. Williams, Kenneth L. and Van Wallach. 1989. Snakes of the world. Vol. 1. Synopsis of snake generic names. Malabar, Florida, Krieger. vii, 234 pp. Page 6 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 Department of Environmental Population and Organismic Biology, University of Colorado, Boulder, Colorado 80309-0334 (HMS); Snow Entomological Museum, University of Kansas, Lawrence, Kansas 66045- 2119 (GWB); Department of Life Science, Northwestern State University, Natchitoches, Louisiana 71497 (KLW); and Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 (VW). Received: 17 June 1991 Accepted: 1 July 1991 Bulletin of the Maryland Herpetological Society Page 7 Volume 28 Number 1 March 1992 DISTRIBUTIONAL AND TAXONOMIC NOTES ON SOME LIZARDS OF THE GENUS SCELOPORUS FROM MEXICO Hobart M. Smith Material recently examined of the genus Sceloporus in the University of Kansas Museum of Natural History (KU), the James Ford Bell Museum of Natural History (JFBM) at the University of Minnesota in Minneapolis, and the University of Colorado Museum (UCM), has revealed several noteworthy range extensions and variations in four Mexican species and subspecies, here reported. Sceloporus bicanthalis Smith Although Smith (1937), in the original description of this taxon, recorded it from “Reyes”, Oaxaca, and later (Smith, 1939) added the locality Mt. Zempoaltepetl in the same state, no records for that state were mapped by Guillette and Smith (1985) in their review of the S. aeneus complex, presumably because of uncertainty of identification. The species is likewise omitted from the herpetofaunal list for the state of Oaxaca in Flores and Gerez (1988: 214). A series of eight specimens of this species (KU 137728-9, 137733-4, 137736-9) from Llano de las Flores (12 mi N Ixtlan de Juarez), 2720-2920 m, Oaxaca, confirms the presence of the species in that state and supports restitution of the other two localities of occurrence. “Reyes” = Papalo Santos Reyes, or Santos Reyes Papalo) is near the extreme northern end of the Sierra de Juarez (about 15 km ENBE of Cuicatlan), and Llano de las Flores is near the southern end of that range, hence there is no question of accuracy of identification of the “Reyes”. It is strange that the species does not occur in the well-collected Sierra deAloapaneca, north of Cd. Oaxaca, just southwest of the Sierra de Juarez, although the latter is faunistically distinctive to a considerable degree. Mt. Zempoaltepetl, on the contrary, is separate from the Sierra de Juarez, but is part of the easternmost Sierra Madre Oriental, to which S. bicanthalis appears to be limited in Oaxaca. All specimens have at least some evidence of dark bars on the throat, as is distinctive of this species; they are especially well developed in the single subadult male (38 mm s-v) which also has dark longitudinal streaks along the sides of the abdomen. The throat bars are clearly defined even in a near-hatchling male (28 mm s-v), but they are dim in all six females, even Page 8 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 the smallest (33 mm s-v). All have 2-2 canthals, and the postrostrals are 1- 1 except for one with 1-2. The dorsals vary from 37 to 41 in all eight specimens; femoral pores 13 -16 in six counts on four specimens; interfem- oral pore scales 0-4 (0,2; 1,2; 4,1). In no respect does the series appear different from other examples of the species. Sceloporus clarki boulengeri Stejneger A single large male (JFBM 2914) from 9 mi S Zacualco, Jalisco, documents the second known locality of occurrence of the species in Jalisco, extending its known range southward about 1 40 km. The specimen is typical of the species, but has 11-11 femoral pores, like the other specimen from Jalisco (Hostotipaquillo; Smith, 1939: 132), but unlike most members of its subspecies, 97 per cent of which have 10 or fewer pores on each side, whereas 96 per cent of S. c. clarki Baird and Girard have 1 1 or more (Smith, loc. cit.). The possibility is suggested that the extreme southern populations of S. clarki may constitute a subspecies distinct from the others. More material is needed to resolve the question. Sceloporus megalepidurus halli Dasmann and Smith Of great interest is the second known specimen (KU 61698) of this subspecies, taken 7.5 mi E (probably SB, off of Hy 175) Tamazulapan, Oaxaca, about 35 airline km SW of the type locality, near San Jose Lachiguiri (Dasmann and Smith, 1974). Unfortunately it is a female, thus not confirm¬ ing one of the subspecies’ diagnostic character states, namely the absence of semeions in males. It is an adult, 49 mm s-v, with 43 dorsals, 14-15 femoral pores, five scales (minimum) between the femoral pore series, basal subcaudals smooth, four postrostrals, canthals 1-1, and supraoculars 4-4, large, in one series. The smooth basal subcaudals (Lynch and Smith, 1965) and small number of dorsals in this subspecies (43-46) differentiate S. m. halli from S. m. megalepidurus Smith (52-63), which is similar in lacking semeions in males. The single row of large supraoculars also distinguishes S. ttl halli from both S. m megalepidurus and S. ttl pictus , although in the original description of the former no special importance was attached to the large supraoculars, presumably interpreted as a nontaxonomic variation. Occurrence of the same state in both specimens, however, as opposed to the regular occurrence of two rows in the other subspecies, suggests that it is taxonomically significant. Sceloporus megalepidurus pictus Smith A single specimen (KU 61697) from 2 mi W Yanhuitlan, 8300 ft, Oaxaca, extends the known range of the subspecies from southern Puebla into the northern part of the state of Oaxaca, a distance of about 140 km Bulletin of the Maryland Herpetological Society Page 9 Volume 28 Number 1 March 1992 (airline) southward from the Tehuacan region (Dasmann and Smith, 1974). Fortunately the specimen is an adult male, 48 mm s-v, with the typical, broadly dark-bordered abdominal semeions, thus differentiating it from both S. m. megalepidurus to the north, S. m. hallito the south. Its scutellation is typical, with 46 dorsals, 14-15 femoral pores, six scales (minimum) between femoral pore series, four postrostrals, canthals 1- 1 , and supraocu- lars in two rows (4-4 in inner row, 3-3 in outer row, latter scales two thirds size of former). Neither of the two other species at present assigned to the megalepidu¬ rus group (S. subpictus Lynch and Smith, 1 965; S. cryptus Smith and Lynch, 1967) can be involved in any intraspecific variation in S. megalepidurus , inasmuch as both have fewer than 38 dorsal scales, whereas 43 is the minimum recorded in any subspecies of S. megalepidurus. Acknowledgments I am much indebted to Drs. Philip J. Regal and James C. Underhill for the privilege of studying JFBM material, to Dr. William E. Duellman and John Simmons for KU material, and Dr. Shi Kuei Wu for UCM material; and to Dr. William M. Lewis for facilities for study. yteratureCited Dasmann, Marlene M. and Hobart M. Smith. 1974. A new sceloporine lizard from Oaxaca, Mexico. Gr. Basin Nat., 34(3): 231-237, figs. 1-4. Flores Villela, Oscar and Patricia Gerez. 1988. Conservacion en Mexico; sintesis sobre vertebrados ter- restres, vegetaciony uso del suelo. Xalapa, Veracruz, Mexico, Inst. Nac. Inv. Rec. Biot, xiv, 302 pp., ill. Guillette, Louis J., Jr. and Hobart M. Smith. 1985. Cryptic species in the Mexican lizard complex, Sceloporus aeneus. Bull. Maryland Herp. Soc., 21(1): 1-15, figs 1-5. Lynch, John D. and Hobart M. Smith. 1965. New or unusual amphibians and reptiles from Oaxaca, Mexico. I. Herpetologica, 21(3): 168-177, figs. 1-4. Smith, Hobart M. 1937. A synopsis of the scalaris group of the lizard genus Sce¬ loporus. Occ. Pap. Mus. Zool. Univ. Michigan, (361): 1-8. Page 10 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 1939. The Mexican and Central .American lizards of the genus Scetopoms. ZooL Ser. Field Mus. Nat. Hist., 26: 1-397, figs. 1-59, pis. 1-31. _ and John D. Lynch. 1 967 . A new cryptic lizard (Iguanidae : Scelopoms) with comments on other reptiles from Oaxaca, Mexico. Herpetologica, 23(1): 18-29, figs. 1-3. *. Environmental Population and Organismic Biology , University of Colorado, Bovlder, Colorado, 80309 0334. Received: 24 June 199 Accepted: 3 July 1991 Bulletin of the Maryland Herpetological Society Page 1 1 Volume 28 Number 1 March 1992 THE NOMENCLATURAL STATUS OF CANTORIA GIRARD, 1858 (REPTILIA: SERPENTES) Hobart M. Smith, Reeve M. Bailey, Kenneth L. Williams and Van Wallach Cantona Girard, 1858, the currently accepted name for a genus of snakes of the Orient and Indo-Australian archipelago, has priority over the name Cantoria Kaup, 1858, for a genus of flatfishes whose currently accepted name is Cynoglossus Hamilton, 1822. As pointed out by Williams and Wallach (1989: 26), the current and long-accepted name Cantoria Girard, 1858, for a genus of snakes occurring from southern Asia southward through the Indo-Australian Archipelago, was proposed in the same year as its homonym Cantoria Kaup, applied to a genus of marine fishes occurring in the eastern Atlantic Ocean of West Africa and in the Indo-West Pacific Ocean from eastern Africa to Japan and Australia. Relative priority of these homonyms is unsettled, although Whitley (1940) assumed that Kaup’s name is the junior one and proposed Cantorusia as a replacement. Girard’s name also has a junior subjective synonym (hence a potential alternative), Hydrodipsas Peters (1859). The most critical fact in this case, relative to nomenclatural stability, is that Cantoria of Girard has long been accepted as the valid name for its genus, whereas Cantoria of Kaup has been regarded as a subjective junior synonym of Cynoglossus Hamilton (1822) in recent decades (e.g. Weber and deBeaufort, 1929: 186, 196; Menon, 1977: 16; Eschmeyer and Bailey, 1990: 74). Therefore no disruption of nomenclature occurs if Kaup’s name is found to postdate Girard’s. Girard’s name appeared at some time after Sept. 29, 1857, when his paper was accepted for publication at a meeting of the Academy of Natural Sciences of Philadelphia (p. 179). The paper was presented earlier, on August 11, 1857 (p. 175). Indeed, 1857 has commonly been accepted as the date of publication (e.g. Smith, 1943; 397; Gyi, 1970: 182), but evidence in Nolan (1913: xi), and the imprinted date for the 1857 volume as a whole (1858), led Williams and Wallach (1989: 26) to accept 1858 as the year of publication. General herpetological practice has been to accept Nolan’s information, largely based on records of dates of receipt of the Academy publications by various eastern U.S. institutions. Unfortunately no records were found of receipt of some separately issued signatures, and obviously not all were promptly acknowledged. Therefore, as indicated by Nolan, Page 12 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 mailing dates (which are the nomenclatural dates of publication) were earlier than the dates given in his compilation, but at least never later. To complicate matters further, "The issue to authors of separate copies of papers from the Proceedings also antedates the publication of the numbers of which they form a part, the record being printed on the covers of the separata but not otherwise preserved.” (Nolan, 1913: vii.) There is, however, no evidence of separate publication of this article. Since Girard’s description of Cantoria appeared in the issue contain¬ ing only signature 13 of the 1857 volume, which ended with the very page ( 182) on which the description was printed, and was presented at the August 1 1 meeting of that year, it seems likely that distribution of that signature may well have occurred in 1857. Indeed, one of us (RMB), in collaboration with R. D. Suttkus, has carefully investigated the publication dates of other articles by Girard, on fishes, that appeared in the Academy Proceedings, and found that commonly an issue appeared within a month or two after the date of the last meeting covered by it, whereas acknowledgment of receipt might not be recorded for eight or more months after that date. There were seven issues of the 1857 volume (vol. 9) (Nolan, 1913: xi). Receipt of the first issue, pp. 1-16, was acknowledged promptly, Feb. 25, 1857, but receipt was acknowledged for only two other issues in the volume: Jan. 7, 1858, for the third issue (pp. 73-100) and May 1, 1858, for the last issue (pp. 183-228). On the basis of that information, given by Nolan, recent workers (e.g. Williams, and Wallach, 1989) have tended to accept 1858 as the date of distribution (publication) of the sixth issue that includes Girard’s article. That view is reinforced by the late receipt of the first issue of the 1 858 volume April 19, 1858, by the American Antiquarian Society, which so promptly acknowledged (Feb. 25, 1857) the first issue of the 1857 volume. Although subjective evidence suggests a strong likelihood that Girard’s paper appeared in late 1857, the objective evidence indicates early 1858, which date we perforce here accept. The first part of vol. 24 of the Archiv fur Naturgeschichte contained Kaup’s article in which he proposed Cantoria. We have found no evidence of publication date for that part, but part two was received at the Philadelphia Academy May 10, 1859, according to a list of “Donations to Library” in vol. II (p. x) of its Proceedings. That evidence, although indirect, strongly indicates that part one was not distributed earlier than late 1858, and may have been as late as early 1859. We thus conclude that, beyond doubt, Girard’s Cantoria 1858 has priority by at least several months over Cantoria Kaup, 1858. Bulletin of the Maryland Herpetological Society Page 13 Volume 28 Number 1 March 1992 Literature Cited Eschmeyer, William N., and Reeve M. Bailey. 1990. Genera of recent fishes. Part 1. Pp. 7-433 in W. N. Esch¬ meyer, Catalog of the genera of recent fishes. California Acad. Sci., San Francisco, v, 697 pp. Girard, Charles. 1858. Descriptions of some new reptiles, collected by the U.S. Exploring Expedition, under the command of Capt. Charles Wilkes, U. S. N. Proc. Acad. Nat. Sci. Philadelphia, 9: 181- 182 (1857). Gyi, Ko Ko. 1970. A revision of colubrid snakes of the subfamily Homalopsi- nae. Univ. Kansas Pubis. Mus. Nat. Hist., 20(2): 47-223. Kaup, Johann Jacob. 1858. Uebersicht der Plagusinae, der funften Subfamilie der Pleu- ronectidae. Arch. Naturg., 24(1): 105-110. Menon, A. G. K. 1977. A systematic monograph of the tongue soles of the genus Cynoglossus Hamilton-Buchanan (Pisces: Cynoglossidae) . Smithsonian Cont. Zool., (238): 1-129. Nolan, Edward J. 1913. An index to the scientific contents of the Journal and Pro¬ ceedings of the Academy of Natural Sciences of Philadel¬ phia, 1812-1912. Philadelphia, Acad. Nat. Sci. xiv, 1419 pp. Peters, Wilhelm C. H. 1859. Nachrichten von Hm. Fedor Jagor, der auf eigne Kosten nach Ostindien und den Philippinen gereist ist und von der Akademie mit Instructionen fur das Sammeln naturwissen- schaftlicher Gegenstande versehen wurde. Monatsb. Akad. Wiss. Berlin, 1859: 269-271, 1 pi. Smith, Malcolm A. 1943. The fauna of British India, Ceylon and Burma. . .Reptilia and Amphibia. Vol. Ill - Serpentes. London, Taylor and Francis, xii, 583 pp. Page 14 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 Weber, Max, and L. F. deBeaufort 1929. The fishes of the Indo-Australian Archipelago. Vol. V. Anacanthini, Allotriognathi, Heterosomata, Beiycomorphi, Percomorphi [in part]. Leiden, Brill, xiv, 458 pp. Whitley, Gilbert P. 1940. The Nomenclator Zoologic us and some new fish names. Australian Naturalist, 10(7): 241-243. Williams, Kenneth L., and Van Wallach. 1989. Snakes of the world. Vol. I. Synopsis of snake generic names. Malabar, Florida, Krieger. vii, 234 pp. Department of Environmental Population and Organismic Biology , University of Colorado, Boulder, Colorado 80309-0334 (HMS): Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109-1079 (RMB); Department of Life Science, Northwestern State University. Natchitoches, Louisiana 71497 (KLW); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 (VW). Received: 6 May 1991 Accepted: 25 May 1991 Bulletin of the Maryland Herpetological Society Page 15 Volume 28 Number 1 March 1992 A SECOND LOCALITY FOR GEOPHIS SALLE! (REPTILIA: SERPENTES) Hobart M. Smith and David Chiszar Seven previously unreported specimens of Geophis salleu in¬ cluding two hatchlings, are from a second known locality for this species: Santa Rosa, nr Lachao, Juquila district, Oaxaca, Mexico. None have a nuchal collar, hence G. laticollaris is resurrected as a valid species from the synonymy of G. sallet The latter species was named for its collector, Auguste Salle, whose patronymic species names should be spelled salleu not sallaeL Although Downs (1967: 171) was aware of eight specimens of Geophis sallei Boulenger when he monographed the genus Geophis , only one precise locality of occurrence was then known for the species: Cafetal Concordia, nearPluma Hidalgo, Oaxaca, Mexico. By virtue of synonymizing G. laticollaris (Smith et al., 1965) with G. salleu an action that we here reject, Downs (1967: 168) added in a footnote the former species’ type locality, 3 mi S Putla, Oaxaca. A series of seven specimens in the University of Colorado Museum (UCM) adds a second locality of record for G. sallei (sensu stricto), and provides a basis for resurrection of G. laticollaris . At the same time we argue for emendation of the specific name from sallaeU as originally proposed by Boulenger (1894: 318), to sallei The specimens are UCM 52609-10, 52616-7 and 52527-9, all from Santa Rosa, near Lachao, district of Juquila, Oaxaca, taken in January, 1971 and July, 1972, by Thomas MacDougall. Lachao is located in the southern foothills of the Sierra Madre del Sur, as is Pluma Hidalgo, about 80 km SE Lachao. The range of the species is presumably more or less continuous between the two localities, since they are topographically similar as well as close geographically. Unfortunately all except UCM 52527-9 are shrivelled and brittle; the exceptions are an adult female (52527), 304 mm s-v, tail 53 mm; an adult male (52528), 365 mm s-v, tail 5 1 mm; and a subadult male (52529) 1 82 mm s-v, tail 31 mm. The others cannot be sexed; two are of medium size, approximately 225 mm (52610) and 260 mm (52609) s-v; and two are essentially hatchlings, about 105 mm (52617) and 125 mm (52616) s-v. Page 16 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 Ventrais and caudals are reliably countable in all, however, the ventrals varying from 123 to 131, the caudals 29-38 (128 and 34-35 respectively, in the sexable female, 126 and 38, 125 and 32, in the two sexable males, respectively). The overlap between the two sexes in both characters is, according to Downs' data (1967: 170), too great to permit accurate determi¬ nation of sex by use of those counts. All specimens are unicolor light to dark brown dorsally, whitish ventrally; the extreme ends of the ventrals are pigmented, and the pigment extends slightly mesad on the anterolateral edges of the ventrals. Cranial scutellation conforms with Downs’ descrip¬ tion, as does the coloration. The most notable feature of pattern is the absence of a light nuchal collar in all specimens, even the juveniles. Downs apparently had no juveniles available to him, but conjectured that they might be collared: on that assumption he synonym ized G. laticollaris , the only known specimen of which is a juvenile, 139 mm s-v, with G. salleu because presence of a collar and larger eyes were the only differences supposedly distinguishing the former from the latter. UCM 52527-9, the only present specimens in which eye size is not distorted, has the eye diameter only very slightly less to veiy slightly more than its distance from the labial border, hence negates any difference in that character, unless it is stepped in geographic variation. But because the holotype of G. laticollaris does have a conspicuous nuchal collar, whereas none of the 15 G. scdlei now known does, even in hatchlings, we regard G. laticoRaris as a distinct taxon. The only locality where it is known to occur, near Putla, is about 120 km from Lachao, considerably farther inland (about 90 km) than either locality known for G. sallei (20-25 km), on northern slopes of the Sierra Madre del Sur, and in a different drainage system. The two taxa could possibly be subspecifically related, but continuity of range seems unlikely. The French collector Auguste Salle obtained important although small herpetological collections in Mexico in the mid- 19th century; his material ultimately went to the Paris and British museums (Kellogg, 1932). Three herpetological patronyms honor him: Anolis sallaei Gunther, 1859 (= Norops sericeus Hallowell); Hylodes sallaei Gunther 1868 (= Eleutherodac - tylus rhodopis (Cope)); and Geophis sallaei Boulenger, 1894. Why Gunther chose to render his eponyms as sallaei rather than sallii or salleii is not apparent; he was presumably attempting to approximate the -ii ending commonly in use at that time for eponyms. However, for another surname ending in -e. Cope, he wrote the name copii (as in Aspidura copii Gunther, 1864, and Leptognathus copii Gunther, 1872). Boulenger apparently was following precedent in adopting the spellings sallaei and copii for patronyms he created in subsequent years. Since all descriptions creating patronyms Bulletin of the Maryjpnd Herpetological Society Page 17 Volume 28 Number 1 March 1992 for Salle and Cope make it clear whom was honored, under Art. 32 (c)(ii) (example) of the Code, all patronyms for both workers should be corrected to sallei and copeu respectively. Ul£mtu£S.£it£d Boulenger, George Albert. 1894. Catalogue of the snakes in the British Museum (Natural History). Vol. II. London, Taylor and Francis, xii, 382 pp., 25 figs, 20 pis. Downs, Floyd Leslie. 1967. Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Misc. Pubis. Mus. Zool. Univ. Michi¬ gan, (131): i-iv, 1-193, figs. 1-23. Kellogg, Remington. 1932. Mexican tailless amphibians in the United States National Museum. Bull. U. S. Natn. Mus., (160): i-iv, 1-224, figs. 1- 24, pi. 1. Smith, Hobart M., John D. Lynch and Ronald Altig. 1965. New and noteworthy herpetozoa from southern Mexico. Chicago Acad. Sci. Nat. Hist. Misc., (180): 1-4, fig. 1, pi. 1. Environmental Population and Organismic Biology, University of Colorado, Boulder, Colorado, 80309-0334 (HMS); and Department of Psychology, University of Colorado, Boulder, Colorado , 80309-0345 (DC). Received: 10 May 1991 Accepted: 25 May 1991 Page 18 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 A MEXICAN GENUS OF TROPIDOPHEINE SNAKES Hobart M. Smith and David Chiszar A dwarf, boa like tropidopheid snake is described, Boella teneihx as a new genus and species from the state of Oaxaca, Mexico, based on a single specimen without exact locality. The productive stem for family-group names based on generic names ending in aphis is op he . The family Tropidopheidae is now a generally recognized group of primitive alethinophidian snakes (Dowling, 1986; McDowell, 1987; Rieppel, 1988), all extant members of which, in McDowell’s (1987) classification, occur in tropical America. Two subfamilies are recognized in the latter work: the Ungaliopheinae, containing two living genera, Exiliboa of Oaxaca, Mexico, and Ungaliophis (Chiapas, Mexico to Ecuador), and the Tropido- pheinae, also containing two extant genera, Tropidophis of the West Indies and northern Southern America, and Trachyboa of Panama, Ecuador and Colombia. An additional genus is here added to the Tropidopheinae, from Oaxaca. Its occurrence in that state along with Exiliboa suggests that separation of their two subfamilies was of great antiquity. The additional genus is represented by a single specimen in a collection pm chased from Thomas B. MacDougall (Smith, 1974) for the University of Colorado Museum (UCM). Unfortunately that particular speci¬ men was one of the few lacking a locality label. However, data on the others provide a clue to its origin. The collection was made from August, 1969, to January, 1973, and numbers about 300 specimens (most under UCM 52376-52670). Localities represented are in eight provinces of the state of Oaxaca, as follows: Ixtlan (Comaltepec, La Esperanza, Llano de las Flores, Vista Hermosa), Juchitan (Palomares), Juquila (Santa Rosa, nr Lachao), Miahuatlan (San Jose, nrLachiguiri), Mixes (Doce de Julio, Ixcuintepec, San Juan Evangelista), Oaxaca (Cerro San Felipe), Putla (San Vicente), and Tehuantepec (numerous localities). The largest series is from Santa Rosa, which lies in one of the least explored regions of Mexico, and it is from there that the snake most likely came. Several other especially notable species are represented in the Santa Rosa series, and all of the other collection sites are relatively well known. Bulletin of the Maryland Herpetological Society Page 19 Volume 28 Number 1 March 1992 Although of uncertain source, limitation of it at least to the state of Oaxaca, and probably to the southern Sierra Madre del Sur, suffice to justify making known this extraordinarily distinct genus and species, even though based on only one specimen. Boella gen. nov. Diagnosis. Small species (total length 580 mm in only known specimen) with a prehensile tail and body strongly compressed laterally; anal spurs present in females (therefore presumably in both sexes); dorsals smooth; scale rows 33-39-21; hyobranchial cornua parallel but separate, short (to 5th ventral) ; ventrals 264; caudals 65, entire; two loreals, one huge, half length of snout, eye to tip of rostral; intemasals and prefrontals paired; nasals in contact behind rostral; kidney not lobulated; pupil vertical; posterior vertebrae keeled. Content. One species, B. tenelia sp.n. Boella tenelia sp.n Holotvpe. UCM 56508, state of Oaxaca, Mexico, collected 1970-71 by Mr. Thomas B. MacDougall. Diagnosis. As given for genus; preoculars 2-2; postoculars 4-4; 4-5 anterior temporals; 11-12 supralabials, 6-6/7 contacting eye; parietals subdivided, anterior pair of scales largest, contacting each other medially, supraocular laterally; nasal paired, nostril in suture between; a small, posteroventral loreal, about size of lower preocular; 6 scales bordering mental groove on each side; scent glands posterior to anal openings; scales of anterior and midbody scale rows largest near middorsum, gradually merging with the much narrower laterals (one-third size of dorsals) except for scales in one or two lowermost rows, which are about the same size as the middorsals; scales in all rows on posterior part of body of about same size, laterals slightly smaller. Description of holotvpe. As given in diagnoses for genus and species, and as shown in accompanying figures; infralabials 12-13; penultimate ventral and first caudal divided; s-v, 491 mm; tail, 89 mm; total length, 580 mm. Light tan above, with numerous, small, dark-brown-edged dorsal blotches, mostly 2-3 scales long and separated by 3-4 scales lengths, crossing most of dorsum; on neck the blotches are fused to form four dark lines that become paired blotches behind neck, these shortly fusing to form median blotches; an irregular series of numerous small dark brown spots. Page 20 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 separated by only one or two ventrals, extending onto lower dorsals. Head more or less uniform brown, somewhat darker than ground color of body; a narrow dark streak 1-2 scales wide from below and behind eye onto neck, continuous with the lower neck streak; supralabials and posterior infrala¬ bials lightly pigmented; rest of infralabials and mental dark; a distinct, short dark line on each side of gular region even with the median (6-11) infrala¬ bials. The right ovary contains 1 0 enlarged follicles, readily distinguishable externally, and the left ovary 8; the largest follicles measure 4-4.5 mm in greatest diameter; only two or three are smaller. We conclude on these bases that the specimen is sexually mature, although it probably is not full grown. The species is thus one of the smallest of the “henophidian” taxa. Comparisons. The five extant genera of tropidopheids, as now known, are remarkably distinct from each other (Bogert, 1968, 1968a). The non-lobulated kidneys and keeled posterior vertebrae of Boella associate it in the Tropidopheinae with Tropidophis and Tr achy boa (McDowell, 1987), but the latter has heavily keeled scales on body, no or a vestigial rostral, nasals separated and single on each side, loreals indistinguishable, no more than 33 midbody scale rows or subcaudals, no more than 152 ventrals, a longer hyoid (to rib 9 or 10), no more than five paramental-groove scales, and no spurs in females. Tropidophis has separated nasals, generally two pairs of prefrontals, no loreals, no more than 29 midbody scale rows or 225 ventrals or 45 subcaudals or five paramental-groove scales, a longer hyoid (at least to rib 7), no spurs in females, and a cylindrical or weakly compressed body. In addition to the lobulated kidneys and flat ventral surfaces of the posterior vertebrae that characterize the Ungaliopheinae (McDowell, 1987) and differ from those features of Boella , both Exiliboa and Ungaliophis have at least the intemasals (and in Ungaliophis also the prefrontals) unpaired, a single loreal on each side, nasals separated, and a cylindrical or weakly compressed body. Ungaliophis also differs from Boella in having the nostril pierced in the anterior of two nasals, no more than 10 supralabials or four paramental-groove scales or 25 midbody scale rows or 258 ventrals or 47 subcaudals, a longer hyoid (to rib 7) and no spurs in females. Exiliboa differs from Boella also in having no more than seven supralabials, three paramental-groove scales, 21 midbody scale rows, 116 ventrals, or 24 subcaudals. The large number of scale rows, ventrals, caudals and paramental- groove scales; two loreals; contact of nasals; and peculiar, greatly reduced Bulletin of the Maryland Herpetological Society Page 21 Volume 28 Number 1 March 199" size 01 the dorsals on sides of body (except posteriorly) ai ; all characters unique to Boella, among tropidopheid genera. Et/mology. Boetiais a diminutive of Boa (Latin, snak ') (Brown, 1954: 490), and tenella is a Latin adjective for “slender.” Remarks. The proper orthography under the Code of family-group names formed from generic names endmg in ophis the Lat in transliteration of a Greek word for “snake,” is subject to different opinions. Following uie unexplained piecedent established by McDcwell (1987) and adopted in several other articles in the same volume. Smith and Preston (1987) argued that the uniquely correct stem of ophis for creating family-group names is ophe-. Contrariwise, Cannatella (1990) maintained that under the Code both stems (op hi- and ophe-) are correct, that the original family-name spellings (all using the ophi- stem), under article 32b (that correct original spellings are to be retained) are therefore to be maintained, and thrt to switch to use of the ophe- stem would, in addition to the preceding, be contrary to the spirit of the Code to maintain stability. Appendix D VII, Table 2. of the Code gives examples as models for determining the stem to be used in formation of (a) derivative and compound words and (b) family-group names. The “grammatical” stem, formed in this case by dropping the final consonant of the nominative form, is, as explained in that table, to be used for the former, whereas the “productive” stem, derived by dropping the genitive singular case ending, is to be used for the latter. The word closest to ophis in orthographical form, given in the table, is /crisis, whose grammatical stem is given as krisi -, its productive stem as krise-. Extrapolating from this example to ophis , the grammatical stem is ophi- (thus correctly yielding “ophidian” and Ophisaurus , for example) and the productive stem ophe-. Article 29b explicitly defers to Appendix D VII for fixation of the standards for determination of the genitive singular stem used in formation of family-group names. Therefore ophe- is the correct stem in present contexts, and ophi- is incorrect. Maintenance of the long-established usage in family-group names of the ophi- stem could be assured by appeal to the International Commission on Zoological Nomenclature. Stability is a most fundamental consideration oi biological nomenclature (hence of the Code), and spelling of names as well as the names themselves fall within the purview of the Commission (Art. 77). Since no confusion whatsover of name application is involved in the present situation, however, appeal for Commission intervention seems unjustified. For these reasons the spellings Tropidopheoidea, Tropidopheidae and Tropidopheinae are utilized here, and a general policy of use of ophe- as the Page 22 Bulletin of the Maryland Herpetologicai Society March 1992 Volume 28 Number 1 proper stem in fo rma tin of all other family-group names derived from generic names ending in ophis is regarded as uniquely correct. Acknowledgments Duane Baxter, Dr. Richard E. Jones and Kristen Lopez aided in study of the visceral anatomy of the holotype. Dr. Shi-KueiWu curated the material and permitted its study, Dr. William M. Lewis provided vital facilities, and Fig. 1 . Dorsal view, head of the holotype of Boella tenellcL Bulletin of the Maryland Herpetological Society Page 23 Volume 28 Number 1 March 1992 Fig. 2. Lateral view, head of the holotype of Boella tenella Page 24 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 Fig. 3. Ventral view, head of the holotype of Boella tenellcu Bulletin of the Maryland HerpetoiogicaS Society Page 25 Volume 28 Number 1 March 1992 Fig. 4. Dorsolateral view of the holotype of Boella tenellcL Page 26 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 Fig. 5. Ventrolateral view of the holotype of Boella tenellcL Bulletin of the Maryland Herpetological Society Page 27 Volume 28 Number 1 March 1992 Literature Cited Bogert, Charles M. 1968. The variations and affinities of the dwarf boas of the genus Ungaliophis. Am. Mus. Novit, (2340): 1-26, figs. 18. 1968a. A new genus and species of dwarf boa from southern Mexico. Am. Mus. Novit., (2354): 1-38, figs. 1-12. Caimatella, David C. 1990. Ancient Greek and ophidian orthography. J. Herp., 24(3): 322-323. Dowling, Hemon G. 1986. Prodromus of a new classification of the Serpentes. II. Arrangement of genera. Privately printed. Pp. 5-22. McDowell, Samuel B. 1987. Systematics. Pp. 3-50, fig. 1*1, in. Richard A. Seigel, Joseph T. Collins and Susan S. Novak, eds., Snakes: ecology and evolutionary biology. Macmillan, New York, xiv, 529 pp., ill. Rieppel, Olivier. 1988. The classification of the Squamata. Pp. 261-293, figs. 7.1- 7.1, in: M. F. Benton, ed.. The phylogeny and classification of the tetrapods. Volume 1: Amphibians, reptiles, birds. Clarendon, Oxford. Smith, Hobart M. 1974. Thomas Baillie MacDougail, naturalist-collector, 1896- 1973. Bull. Maryland Herp. Sex*., 10(3): 89-94, figs. 1-2. _____ and Michael J. Preston. 1987. The stem for formation of family-group names from the Greek word ophis. Bull. Maiyiand Herp. Soc., 23(3): 128- 129. Department of Environmental Population and Organismic Biology , University of Colorado , Boulder, Colorado » 80309 0334 (HMS); and Department of Psychology , University of Colorado , Boulder ; Colorado , 80309 0345 (DC). Received: 5 November 1991 Accepted: 21 November 1991 Page 28 Bulletin of the Maryland Herpetological Society It's what Volume 28 Number 1 March 1992 ‘NT.'WS & ‘Kotes Bulletin of the Maryland Herpetological Society Page 29 Volume 28 Number 1 March 1992 O^OTIS: Lecture and Field Trip Schedule Maryland Natural History Society 2643 North Charles Street, Baltimore, Md. (Phone 301-235-6116) All lectures will be at the Society’s Headquarters at 2643 North Charles Street, Baltimore, Md. For more information call the office at 301-235-6116 between 9:30am and 4pm on Wednesdays. February 12 - Wednesday, 8:15pm - Monthly Lecture "The Maryland Natural Heritage Program: Identifying and Managing Our Rare Species." Janet McKegg - Maryland Department of Natural Resources February IS - Saturday, all day - Field Trip Join us for a trip to the Calvert Marine Museum at Solomons. This remarkable new facility specializes in the maritime history of the Chesapeake. In addition to numerous cultural exhibits (an extensive collection of authentic bay boats, an entire lighthouse, historic photographs, etc.), they have a superb collection of Miocene fossils. We will have a chance to wander through the exhibit areas, visit the fossil lab and preview the fossil exhibits that are presently under construction. We will carpool from the metro area and meet at the museum. Bring you own lunch or plan to visit one of the nearby restaurants. Museum entry fee of $3.00. Contact Butch Norden for information at 410-974-3589 or leave a message at 235-6116. March 5 - Thursday, 8:15pm - Special Lecture "A Naturalist’s Ramblings through the Baja Peninsula." David S. Lee - North Carolina State Museum of Natural Science. Mr. Lee, a former Baltimore resident and long time member of the Maryland Natural History Society, is well known in the field of herpetology. March 25 - Wednesday, 8:15pm - Monthly Lecture "Experiences With Natural History Museums from California to Delaware." Glen Ives - Director, Delaware Museum of Natural History ***NOTE***This exciting lecture coincides with our 63ed annual meeting and yearly social. Nonmembers and members alike are invited to join us for wine and cheese as we celebrate the long history of the Maryland Natural History Society. April 11 - Saturday, All Day - Field Trip We are off to Devonian and Pennsylvanian fossil localities in western Maryland. This one day trip will take us to several interesting sites that usually yield a wide variety of invertebrate and plant fossils 270 to 400 million years in age. We will carpool from the metro area and meet at the Park and Dine Restaurant in Hancock for breakfast. Contact Butch Norden for additional details at 410-974-3589 or leave a message at the Society headquarters. Bring your lunch and rock hammer. April 15 - Wednesday, 8:15pm - Monthly Lecture "The Role of the U.S. Fish and Wildlife Service in the Conservation of Rare and Endangered Insects." Judy Jacobs - U.S. Fish and Wildlife Service May 16 - Saturday, All Day - Field Trip Meet at the Smithsonian in Washington to see and discuss Maryland dinosaur fossils. After lunch we will visit some nearby sites where fossils have been collected. Size of group will be limited so contact Butch Norden to make arrangements, 974-3589. We will meet in front of the Natural History Museum at 10am (parking along mall is prohibited before ten, but if you arrive between 9:30 and 9:45 you can park and sit in your car). Lunch can be purchased at the Museum (ok, but overpriced) or bring your own and we can eat on the mall by the Triceratops. -continued- Page 30 Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 ‘JtE'WS & ‘HpWES May 13 - Wednesday, 8:15pm - Monthly Lecture "Recent Changes in the Definition of Species Based upon Research on Hybrid Ferns." - Donnell Redman SPRING NATURE WALKS AT CYLBURN THURSDAYS - 9:00 AM Meet at Cylbum Mansion House, 4915 Greenspring Avenue NOTE: The walks are canceled if it is raining. Special help for beginning birders. April 2 Winter & Residential Birds, Wildflowers. Leaders: Jean Worthley and Christine Bowen. April 9 Wildflowers, Early Nesting Birds & Possible Migrating Birds. Leaders: Jean Worthley and Christine Bowen April 16 Wildflowers & Migrating Birds. Leaders: Jean Worthley and Christine Bowen. April 23 Migrating Birds and Wildflowers. Leaders: Jean Worthley, Christine Bowen and Erana Lubbert. SPRING - ALL COUNTY WALKS Meet at the rear of Howard Johnson Restaurant, Dulaney Valley Road & Fairmont Avenue, Towson. We will leave promptly at 8:00 AM. BRING LUNCH. For additional information on meeting place - phone Erana Lubbert 301-339-6133. NOTE: All walks canceled if it is raining. April 30 Susquehanna State Park - Harford County (All Day) For Gleason’s Trillium by the thousands, other spring flowers and migrating birds. Leaders: Rodney Jones and Erana Lubbert May 7 Monkton - Dr. Dandy’s Fissure Estate (All Day) Bobolinks & Yellow Orchids and much, much more. Leaders: William Gray and Erana Lubbert May 14 Woodstock (All Day) To Patapsco River for Cerulean Yellow Throated plus other resident & migrating birds. Wildflowers also. Leaders: William Gray and Eiana Lubbert May 2 1 Gunpowder State Park Long but easy walking trail. Migrating birds & spring wildflowers. Leaders: William Gray and Erana Lubbert LEADERS Christine Bowen William Gray Rodney Jones 301-859-8200 301-730-5655 301-486-3442 Erana Lubbert Jean Worthley 301-339-6333 301-374-5370 Bulletin of the Maryland Herpetological Society Page Volume 28 Number 1 March 1992 9&TES: Prepublication Announcement A Revised Checklist with Distribution Maps of the Turtles of the World by John B. Iverson Publication date: April 1992 This is a substantial revision of Iverson's 1986 Turtle Checklist. At approximately 370 pages, it is 33% longer than the earlier version, and is now the most comprehensive guide available to the scientific nomenclature of the turties of die woilu. The Checklist includes updated distribution dot maps (nearly 270 total) for all species, and information on the original citations, type specimens, type localities, distribution, and pertinent literature for all recognized species AND subspecies. Dichotomous keys modified from the scientific literature are also included for identification down to the species level. Phylogenetic relationships are also illustrated for most taxa. This edition is currently available only in soft cover. Cost is $25.00 prepaid (including surface shipping; foreign airmail shipping is $10.00 extra). All payments must be in U.S. dollars, drawn on U.S. banks, or on an international money order. CaRETTOCHELYIDaE Address order to: John B. Iverson Dept, of Biology Earlham College Richmond IN 47374 USA Enclosed is my check for $ _ . Please send me _ copies of the Checklist. Name: _ _ Address: - _ Page Bulletin of the Maryland Herpetological Society Volume 28 Number 1 March 1992 y&WS & ‘N&FES FeDruar-y 9, 1992 ASSISTANT NEEDED FOR WILDLIFE STUDY PROJECT: THE DISTRIBUTION AND ABUNDANCE OF HERPETOFAUNA AMONG HABITAT TYPES AT REMINGTON FARMS, CHESTERTOWN, MARYLAND. RESPONSIBILITIES: The successful applicant will assist a graduate student in conducting a community level study of amphibians and reptiles inhabiting different habitat types on Remington Farms. Duties will include constructing and installing drift fences and live traps, collecting, measuring, weighing, marking, and releasing amphibians and reptiles; and measuring and recording various habitat parameters. QUALIFICATIONS Experience in wildlife ecology, zoology, biology, or herpetology would be helpful, though not necessary for this position. A keen interest in amphibians and reptiles is desirable. STARTING AND ENDING DATE The assistant will be required from mid-February through May, 1 992. A part-time schedule may be • • possible for a longer period of time if so desired. SALARY $5.00/hr plus housing on the farms if needed. Interested applicants should contact Rod McLeod or Mark Conner at: Remington Farms Rte 2, Box 660 Chester town, MD 21620 (410) 778-3223 or (410) 778-8400 Bulletin of the Maryland Herpetological Society Page Volume 28 Number 1 March 1992 % (OTES: Second World Congress of Herpetology 29 November 1993 - 6 January 1994 For information please contact: Michael J. Tyler, Department of Zoology, University of Adelaide, Box 498, GPO Adelaide S.A. 5001, Australia. FAX: 61-8-223-5817 TERRY WILKINS' WORLD OF REPTILES: A series of videotapes designed for the novice and professional reptile enthusiast. Each 65 minute program discusses the individual species history, habitat, acquisition, care, feeding, breeding, cage design and construction, medical treatment and common asked questions. Each tape is $38.42 ($34.95 plus tax and shipping). Tapes available include: Burmese Python, Ball Python, Reticulated Python, Rat Snake, King Snake. Contact: TERRY WILKINS, P.O. BOX 170, PICKERINGTON, OHIO 43147. 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The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc. 2643 North Charles Street Baltimore, Maryland 21218 US ISSN: 0025-4231 BULLETIN OF THE ^Rarylanb f)ecpetologtcaI 0oriety DEPARTMENT OF HERPETOLOGY THE NATURAL HISTORY SOCIETY OF MARYLAND, INC. MdHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League 30 JUNE 1992 VOLUME 28 NUMBER 2 BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY Volume 28 Number 2 June 1992 CONTENTS NOMENCLATURE FIXATION OF SCELOPORUS FORMOSUS (REPTILIA: SAURLA) AND THE STATUS OF S. SALVINI Hobart M. Smith and Gonzalo Perez-Higareda . . . 31 SCELOPORUS HORRIDUS AND S. SPINOSUS (REPTILIA: SAURLA) ARE SEPARATE SPECIES Hobart M. Smith and David Chiszar . .....44 A REDESCRIPTION OF THE LARVAE OF RANA MONTEZUMAE (ANURA: RANIADAE) John K, Korky and Robert G. Webb . . . . 53 AN ALBINO THAMNOPHIS SIRTAUS FROM FAIRFIELD COUNTY, SOUTH CAROLINA Chuck Smith and Kyle Schuett . . . . . 65 The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc. 2643 North Charles Street Baltimore, Maryland 21218 BULLETIN OF THE mbbs Volume 28 Number 2 June 1992 The Maryland Herpefological Society Department of Herpetology, Natural History Society of Maryland, Inc. Executive Editor... . Herbert S. Harris, Jr. Steering Committee Frank Groves Jerry D. Hardy, Jr. Herbert S. Harris, Jr. Library of Congress Catalog Card Number: 76-93458 Membership Rates Membership in the Maryland Herpetological Society is $16.00 per year. Foreign $20. 00 /year. Make all checks payable to the Natural History Society of Maryland, Inc. Meetings The third Wednesday of each month, 8:15 p.m. at the Natural History Society of Maryland (except May— August, third Saturday of each month, 8:00 a.m.). The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 p.m. Volume 28 Number 2 June 1992 NOMENCIATURAI FIXATION OF SCELOPORUS FORMOSUS (REPTSLIA: SAUR1A) AND THE STATUS OF 8. SALVINI Hobart M. Smith and Gonzalo Perez-Higareda A lectotype is designated for Sceloporus formosus, and its type locality restricted to the vicinity of Xalapa, Veracruz. One lec to para- type represents S. salvini and presumably also is from the vicinity of Xalapa. The latter species occurs in mountains southeastward to southwestern Guatemala, and in the isolated Los Tuxtlas Moun¬ tains, as well as southwestward from the Isthmus of Tehuantepec at least to the Pluma Hidalgo area. S. intemasalis is a junior synonym. The species is unique in having highly fragmented intemasal scales and the fro n to nasals separated from each other. No subspecies are recognized. As recorded by Taylor (1969: v), Wiegmann’s (1834: 50, pi. 7 fig. 2) Sceloporus fomiosus is documented nomenclatu rally by not only the original description but also by the specimens under six numbers, 632-637, in the Zoologisches Museum in Berlin. Taylor stated that those numbers pertained to six specimens, but actually no. 633 was applied to two specimens; the others were applied to one each. Inasmuch as the proper allocation of the name Sceloporus formosus is in some doubt, due to the multiplicity of the taxa in its group. Dr. Gunther Peters, then (1978) curator of herpetology in the Berlin Museum, kindly loaned to HMS the entire series of seven syntypes of that species. The press of other duties has prevented study of that material until the present time. In the interim, several office and laboratory moves and the departure of an intended collaborator conspired to misplacement of the two specimens under no. 633; they cannot now be found. Dr. Rainer Gunther has neverthe- lesss kindly extended the loan to permit completion of the study; all five remaining specimens have now been returned to the Berlin Museum. Of the five specimens, four (634-7) are clearly representative of the species to which the name has been applied, as in Smith (1939). The fifth, no. 632, represents the species now known as S. salvini The localities from which these syntypes came are unknown. How¬ ever, Stresemann (1954) outlined the travels in Mexico of Ferdinand Deppe, who undoubtedly was solely responsible for collecting the material, al¬ though Count von Sack, a wealthy nobleman, is recorded in the Berlin Bulletin of the Maryland Herpetological Society Page 31 Volume 28 Number 2 June 1992 Museum catalog as a co-collector of the syntypes. Actually Deppe seems not to have travelled with the Count in Mexico at all; they debarked at Alvarado, Veracruz, in December, 1824, and Deppe remained in the count’s employ until May, 1825. The dual attributions of collectors for these syntypes suggests that they may all have been collected in early 1825, when they were at least nominally together. Certainly during that time Deppe is known to have travelled rather extensively in central Veracruz, whence it is presumed that the specimens came. He is known to have gone to Mexico City in early 1825 via Xalapa, not by the Orizaba route, which he would have taken had he gone through Acultzingo, to which the type locality was restricted by Smith and Taylor (1950: 346). There is no record that he ever took the Acultzingo- Orizaba route to Veracruz, hence we here revise the type locality for Scelopoms formosus to Xalapa, Veracruz, where it is certain that Deppe visited and that S. formosus occurs. Variation among the four syntypes of that species includes what is known for that area; at least three of them seem typical for that region. It is highly likely that the syntype representing S. salvini also came from the Xalapa area; the species is already known from there, and indeed hitherto only there in the state of Veracruz. Scelopoms salvini Gunther The single syntype referred to this species (ZMB 632) is an adult female 88 mm s-v, tail regenerated. Nasals separated from rostral by one scale, narrowly contacting lorilabials on one side, narrowly separated on the other. Intemasals fragmented, four in a transverse row between nasals, posterior to the four postrostrals, and six in the following row, followed by three scales between the anteromedial edges of the lateral frontonasals; a scale separating the latter on each side from the small median frontonasal; rear apex of latter split off as a very small, separate scale; prefrontals very narrowly in contact medially; frontal ridges prominent; supra oculars in two rows, but the lateral scales (3-4) 1/3 to 1/4 size of the adjacent four largest medial scales (4-5); frontal divided, narrowly contacting interparietal, sepa¬ rating the two frontoparietals; latter contacting one rear supraocular on one side for about 1/2 the latter’s medial border. Two canthals, the anterior situated above the canthal ridge one side; head scales essentially smooth, but most with sensoiy pits, usually along their free posterior edge. Dorsals 37 interparietal to base of tail; femoral pores 12-12, the two series separated by 13 scales. Specimen discolored, slightly distorted, slightly soft in abdomen; outer scales shed on anterior third of trunk, where the underlying scales appear blue -black; a narrow black collar appears to be present in front of the forelegs involving two or three scale rows near Page 32 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 mi dime, complete dorsally (a possible midline interruption may exist) and on sides, but not evident vent rally; on middle and rear part of dorsum of trunk, the scales have not been shed, and reveal irregular, narrow (mostly one scale width) , broken crossbands or transverse series of spots extending onto sides and continued on tail as a paired series of spots. This syntype is identified as S. salvini rather than S. formas us because of the numerous intemasals, separation of the median from the lateral frontonasals, and the relatively few femoral pores. In S.formosus the femoral pores are never fewer than 13, the intemasals never so reduced in size, and the median frontonasal always contacts the lateral frontonasals. Much more critical is a comparison with S. intemasalis Smith and Bumzahem (1955). Stuart (1971) vastly expanded the concept of range of that species, to include southern Veracruz (vicinity of San Andres Tuxtla) and montane southwestern Guatemala, focussing primarily upon the extensive fragmentation of the scales in the intemasal and median fronto¬ nasal areas, accompanied by a relatively small number of femoral pores (2 1 - 27 total), compared with S.formosus , and moderate-sized dorsals (35-40). However, those features occur also in S. salvini as noted by Smith (1939: 55- 56). Stuart (1971) dismissed that species from consideration in the complex with fragmented intemasals because it is described as having “two well- developed rows of supraoculars”, but that feature is subject to considerable variation at least west of the Isthmus of Tehuantepec. Since no appraisal of the fragmented-intemasal species, referred to as S. intemasalis by Stuart ( 197 1), as it occurs in Mexico, has appeared since Stuart’s revision, a survey was made of all readily available material from west of the Isthmus of Tehuantepec. It has become apparent that the names S. salvini and S. intemasalis are based on the same species, and although considerable geographic variation exists, it does not appear consistent enough to warrant recognition of subspecies at the present time. In Mexico, the species is very readily and infallibly distinguished from all members of the formosus group by separation of the medial and lateral frontonasals from each other. One, two or all of those scales are commonly reduced in size, in conjunction with fragmentation of the inter¬ nasal scales which in extreme cases are reduced to rounded granules, but in others remain angular although irregular and of moderate size. S. formosus , S. stejnegeri and S. tannerl all members of the formosus group occurring in Oaxaca and Guerrero (S. stejnegeri very likely extends into western Oaxaca, although not yet recorded there), all have the three frohto- nasals large and in contact with no or very rare exception, and the intemasals are not particularly fragmented. Stuart (1971) has confirmed the nearly perfect constancy of the frontonasal character in the species? ih the S r- ( \n^h /; o. v Bulletin of the Maryland Herpetological Society Page 33 Volume 28 Number 2 June 1992 Guatemala, Chiapas, Veracruz and Oaxaca material he examined (two exceptions in 39 specimens from Chiapas, two in 40 from Guatemala, none in ten from Oaxaca and Veracruz). Material we have examined extends the known range of S. salvini somewhat, and fleshes out the range in other parts, as follows. VERACRUZ: Xalapa (MCZ 46935, 136000-5); presumably Xalapa area (ZMB 632); 20 km ENE Jesus Carranza, 200 ft (KU 27511-2); 25 km SE Jesus Carranza (KU 27513); Estacion de Biologia Tropical Los Tuxtlas, 150 m (UCM field 160); Balzapote, Los Tuxtlas area (UCM 51890); 2.5 mi N La Palma, Los Tuxtlas area (UTA 3097); Arroyo Claro, Sierra de Santa Marta, Los Tuxtlas area, 1000 m (UCM field 323-7). OAXACA: West of Figueroa, 5000 ft (a few km NE Pochutla) (UCM 39835-8); Metates, N slope Sierra Juarez, 17.7 km S Valle Nacional (UTA 11785, 24008, 11783-4); San Pedro Yetla, 7 km S Valle Nacional (KU 87372); 2 mi E Tollocito (KU 40146); 2 mi S Tollocito (KU 40147); Vista Hermosa (UCM 39839, 49443). CHIAPAS: Linda Vista, 2 km NW Pueblo Nuevo Solistahuacan, 1675 m (KU 59694): Cerro Ovando (UCM 39661). The species has a considerable vertical range, from 61m (20 km ENE Jesus Carranza, Veracruz) to 2500 m (Cerro Paxtal, Chiapas). Its geographic range extends from central Veracruz through northern Oaxaca and across the Isthmus of Tehuantepec onto the uplands of Chiapas to extreme western Guatemala, and along the Sierra Madre del Sur of Oaxaca at least as far southwest as Pochutla. Distribution over much of this range appears to be rather spotty, with local variational tendencies whose extremes are usually matched in an irregular pattern elsewhere. The fewest femoral pores (10) occurs most frequently in the Balzapote and Sierra de Juarez series, but the highest number ( 1 5) occurs in the Sierra de Santa Marta series, very near Balzapote, and in the Figueroa series. The Jalap a series has the highest mean dorsal scale count (40.5), with a range in 8 specimens of 37-44. A high of 40 occurs in the Figueroa and Arroyo Claro series, and all other counts are lower, 35- 39. The total range of dorsals, 35-44, is not exceptional. The supraoculars are highly variable, from one row of large scales in some specimens of widely scattered occurrence, especially common in the Los Tuxtlas area, to two rows, the outer of which consists of 1-4 very small to large (one half to two thirds size of inner supraoculars) scales; the Jalapa series has the largest outer supraoculars, but they vary from small to large and are matched in every other sizeable series. The black collar is most prominent, broadest (2-5 scales) and most frequently complete, or nearly so, in the Jalapa series, but it is closely matched in the Sierra de Juarez series, and to varying degrees in others; a Page 34 Bulletin of the Maryland Herpetoiogical Society Volume 28 Number 2 June 1992 width of three rows is common everywhere, and drops to two frequently. The middorsal interruption varies from one to several scales in various parts of the range. In general the collar width appears to diminish toward Guate¬ mala, and the size of the medial interruption appears similarly to increase. These erratic variations appear to preclude recognition of any geographic races. All have two canthals and separation of the nasal from the iorilabials, with one exception each. The supraoeulars contact the parietal in none, conforming in these respects with Stuart’s (1971) characterization of S. tntemascdts (=S. salving. The largest specimen examined was a female 100 mm s-v (UCM Held no. 326), and the largest male measured 94 mm s- v (UCM field no. 323). The variation occurring in the representative samples examined of S. solvtni leaves no question of assignment of ZMB 632 to that species. Inasmuch as a lectotype is designated in the following pages for S, formosus , ZMB 632 becomes a lectoparatype and is no longer to be considered a syntype fide the 1985 Code. Scelopoms formosus Wiegmann Lectotvpe. ZMB 635, an adult male, the largest of the syntype series of this species. Paralectotvpes ZMB 634, 636 and 637, of those presently at hand; no. 637 is a female, the other two males, and all are adults. The two specimens at present misplaced, under the number ZMB 633, are likewise lect op ara types. None is to be considered a syntype in the future. Description of the Lectotvpe (see Figs. 1-3). Specimen somewhat soft abdominally, but not discolored; s-v length 83 mm, tail 105 mm; dorsals 36; femoral pores 17-18; scales between pore series 6. Uniform bluegreen above and on sides, except for a black collar in front of arms, complete ventrally but widely broken dorsally by 8-9 scales. Other characters as depicted in Figs. 1-3. The main reasons for selecting this specimen as lectotype are not only that it is the largest of the series, is the least distorted and exemplifies very well the distinctive characteristics of the species with which the name has always been associated, but it appears to be the very one illustrated in the original description (Wiegmann, 1834; pi. 7, fig. 2, correctly identified in the text, p. 50, but erroneously stated in the legend on the plate as representing S. torquatus) . The dark borders of the abdominal semeion, and the black ventral collar on the neck, are shown in the figure exactly as in the lectotype, and the hind legs are positioned exactly the same way. The only notable difference is that in the lectotype it is the right arm that is extended Bulletin of the Maryland HerpetoSogical Society Page 35 Volume 28 Number 2 June 1992 forward, the left directed posteriorly, rather than the reverse as shown in the figure; presumably an artistic license to switch the limbs was exercised for esthetic reasons. A cause for some concern in selecting ZMB 635 as lectotype is the relatively small size of the supraocuiars of the inner row, and the relatively large size of those in the outer row. Normally the scales of the inner row are 3-4 times the size of those in the outer row, and in some specimens there is but one row. The variation in size of the supraocuiars may well be geographi¬ cally correlated, although not wholly so. Primarily because all of the Deppe specimens of this species collected while associated with Count von Sack presumably came from the Xalapa area (to which we have here restricted the type locality), as indicated by the itinerary deciphered by Stresemann (1954), we conclude that the small supraocuiars of the lectotype represent merely a variant of the Xalapa population. The three lectoparatypes of this species have typically large supraocuiars in the inner row. Lectoparatypes. The three lectoparatypes referred to S. formosus , ZMB 634, 636 and 637, have the following characters, respectively; male, male, female; dorsals 34, 32, 36; largest lateral supraocular 1/4 - 1/5, 1-6, 1/6 size of largest medial supraocular; 3-3 lateral supraocuiars in all; intemasals 2-2, 2-1, 2-2; femoral pores 16-16, 15-16, 14-15; scales between pore series 7, 7, 4; s-v length 77 mm, 60 mm, 73 mm. The males are blue- green above, not discolored, have a dark slash in front of arm insertions continuous across throat in 634, narrowly interrupted midventraily in 636, ends tapering to a point dorsally on each side, widely (5, 7 scale widths, respectively) separated. The abdominal semeions are prominent, reaching from axilla to groin, with a black medial border about three scales wide at midbody, much wider in axilla and groin, separated at midbody by 3-4 scale rows, not encroaching onto chest or sacral regions. Throat bluish, lighter anteriorly in 636. The single female appears dorsally much like the males, due to loss of most outer layers of the scales; ventral surfaces unmarked; black patch in front of arm insertions small, not extending onto ventral surface. Acknowledgments We are greatly indebted to Drs. Gunther Peters and Rainer Gunther for the privilege of studying the former syntypes of S. formosus: HMS abjectly apologizes for the hopefully temporary loss of two of those specimens. We are also grateful to Dr. William M. Lewis, Jr., for provision of facilities for study; and to Dr. Wm. E. Duellman, John E. Simmons, Dr. Jonathan A. Campbell, Jose P. Rosado and Dr. Shi Kuei Wu for the privilege of studying material in their museums. Page 36 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Fig. 1 . Dorsal view of lectolype, ZMB 635, of Sceloporus formosus , 83 mm s-v. Bulletin of the Maryland Herpetological Society Page 37 Volume 28 Number 2 June 1992 Fig. 2. Ventral view, same specimen as in Fig. 1 . Page 38 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Fig. 3. Dorsal head scales, same specimen as in Fig. 1. Bulletin of the Maryland Herpetological Society Page 39 Volume 28 Number 2 June 1992 Fig. 4. Dorsal head scales of a paralectotype (ZMB 637) of Sceloporus formosus , showing the typically large supraoculars of the species. Page 40 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Fig. 5. Dorsal head scales of the paralectotype (ZMB 632) of Sceloporusformosus that actually represents Sceloporus salviriL Bulletin of the Maryland Herpetologicai Society Page 41 Volume 28 Number 2 June 1992 Fig. 6. Dorsal head scales of Sceloporus salvinu field no. 1 60, from the Los Tuxtlas area, southern Veracruz, showing the highly fragmented intemasals characteristic of the species, and notably more fragmented than in ZMB 632 (Fig. 5). Page 42 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Literature Cited Fugler, Charles M. and Robert G. Webb. 1957. Some noteworthy reptiles and amphibians from the states of Oaxaca and Veracruz. Herpetologica, 13(2): 103-108. Smith, Hobart M. 1939. The Mexican and Central American lizards of the genus Sceloporus. Zool. Ser. Field Mus. Nat. Hist., 26: 1-397. ______ and Carlos B. Burnzahem. 1955. The identity of the Trans-Isthmic Mexican populations of the Malachite tree lizard ( Sceloporus malachiticus Cope). Herpetologica, 11 (2): 118-120. _____ and Edward H. Taylor. 1950. Type localities of Mexican reptiles and amphibians. Univ. Kansas Sci. Bull., 33(8): 313-380. Stresemann, Erwin. 1954. Ferdinand Deppe's travels in Mexico, 1824-1829. Condor, 56(2): 86-92. Stuart, Laurence C. 1971. Comments on the malachite Sceloporus (Reptilia: Sauria: Iguanidae) of southern Mexico and Guatemala. Herpetol¬ ogica, 27(3): 235-259. Taylor, Edward H. 1969. Wiegmann and the herpetology of Mexico. Soc. Study Amphs. Repts., Facs. Repr. Herp., (23): iii-vi. Wiegmann, Arend F. A. 1 834. Herpetologica mexicana seu descript io amphibiorum Novae Hispaniae. Pars prima. Saurorum species. Berlin, Luderitz. vi, 54 pp. Department of Environmental Population and Organismic Biology, Univer¬ sity of Colorado, Boulder, Colorado, 80309-0334 (HMS); and Estacian de Biologia Tropical Los Tuxtlas, Apartado Postal 51, Catemaco, Veracruz (GPH). Received: 3 August 1991 Accepted: 20 August 1991 Bulletin of the Maryland Herpetoiogical Society Page 43 Volume 28 Number 2 June 1992 SCELOPORUS HORRIDUS AND S. SPINOSUS (REPTIUA: SAURIA) ARE SEPARATE SPECIES Hobart M. Smith and David Chiszar Sceloporus horridus (with its three subspecies, S. h. horridus. S. h . oligoporus and S. h. albiventris) and S. spinosus (with its three subspecies, S. s. spinosus , S. s. caendeopunctatus , and S. s. apicalis) are separate species, occurring sympatrically from south- western and southern Puebla to north-central Oaxaca. Intergrada¬ tion is documented between S. s. apicalis and both S. s. spinosus and S. s. caeruleopunctatus, but seemingly does not occur between the latter two subspecies, contrary to the previous concept. Critical material examined from southern Puebla and Oaxaca in the collections of eight museums (AMNH, CAS, FMNH, KU, UCM, tJIMNH, UMMZ, UTA; see Leviton et al., 1980) has made it evident that, contrary to the proposition by Boyer et al. (1983), Sceloporus horridus Wiegmarm and S. spinosus Wiegmann are allospecific, not conspecific, and that S. s. apicalis Smith and Smith, not S. s. caeruleopunctatus Smith, intergrades with S. s. spinosus. A. Allospecificity of S. horridus and S. spinosus Proposal by Boyer et al. (1983) that S. horridus and S. spinosus are conspecific was predicated on intermediacy between the usual character- states of the two oculofrontoparietal series of scales in the two species as seen in a series of nine specimens from near Tlaxiaco, Oaxaca, a locality between the known ranges of the two species. No localities of sympatry were then known between the extensively parapatric ranges of the species. However, material now available documents sympatry in at least two locali¬ ties, and an overlap of their ranges of about 150 km from southern Puebla to central northern Oaxaca (Fig. 1). Localities and areas of note are as follows. 1 . San Diego. Puebla. This locality, about 2 km SB Tehuacan, is represented by six specimens of S. spinosus (UMMZ 195757 (1 spec.) and 88617 (5)) and 15 of S. horridus (UMMZ 88615 (10 spec.) and 88616 (3), and UIMNH 48943, 48945). The six S. spinosus have 7-10 (x 8.7) femoral pores, the 15 S. horridus 3-5 (x 4.2) on each thigh, and this is the most reliable difference between the two species. One supraocular is in contact with the Page 44 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 median head scales on both sides of one S. spinosus, but in all S. horridus except on one side of two. The close approximation of other locality records throughout much of the surrounding area, for both species, suggest widespread sympatry in that region. 2. 0,5 and 2 mi S. Guelatao. Oaxaca. This is the southeasternmost known locality of occurrence ofS. horridus , about 56kmNECd. Oaxaca. Five adult specimens (UIMNH 60452-3, UCM 38313-5) appear typical, with 29- 30 dorsals, 3-5 femoral pores and 4-4 supraoculars of which one is in contact with the median head scales. One S. spinosus (UCM 38364), a subadult male, has 30 dorsals, 7-7 femoral pores, and 4-4 supraoculars; the rear supraocular is in contact with the median head scales on each side. The latter condition is typical of S. horridus , but does occur in a small percentage of S. spinosus. Never do seven femoral pores occur in S. horridus , however, so far as is now known, hence no. 38364 is unquestionably referable to S. spinosus. 3. Other S. horridus from Oaxaca. A typical specimen (CAS 87311) from 2 mi SW Ixtlan de Juarez, 5400 ft (28 dorsals, 5-5 femoral pores, 4-4 supraoculars, rear one contacting median head scales), confirms the occurrence noted previously near Guelatao; the collecting sites may well be exactly the same. On the road southeast from Tehuacan, a locality 2 mi S Teotitlan, just across the Oaxaca border, is represented by two specimens (UCM 38336-7), and, some 30 km farther southeast, a locality 2 mi S Tecomavaca is represented by one (UCM 38335). To the southwest from Tehuacan, the species has been taken 59 km SW Acatepec, Puebla (near Cuyotepeji) (UTA 6386), at El Espinal, 1 1.6 km S jet Hys 126 and 190, on 190 (UTA 25799), and 2 mi NW Tamazulapan (UTA 33807, 48997). These seven specimens conform with previously established limits of the species, having 29-32 dorsals (x = 30), 3-6 femoral pores (x = 4.5), 4-4 supraoculars except for one (33807) with 5-5, and one supraocular on each side contacting the median head scales except for one (38336) with none. Additional records in the literature (Smith, 1939) for S. horridus in Oaxaca are for Chazumba (just across the Puebla border southwest of Tehuacan), Huajuapan de Leon (on the same road as the preceding, at its junction with the Puebla-Oaxaca highway), and Cuicatlan (hallway between Tehuacan and Oaxaca, to the southeast of Tehuacan). 4. Other S. spinosus from within the range of S. horridus. Although S. spinosus ranges widely in south-central Oaxaca, we are aware of but two localities of record from within the range of S. horridus in Oaxaca. One (near Bulletin of the Maryland Herpetological Society Page 45 Volume 28 Number 2 June 1992 Guelatao) has already been mentioned. The other is at El Tejocote (UTA 8602, 11804-5). These specimens appear to be typical of S. s. spinosus, having 10-10 femoral pores in all, 27-28 dorsals and 4-4 supraoculars; the supraoculars are separated from the median head scales on both sides of one, one side of another. In Puebla, Cope (1885: 379) reported S. spinosus from Tlapanala, about 13 mi N Izucar de Matamoros. Inasmuch as Cope provided a synopsis of the genus in the same work, characterizing S. spinosus with 10 femoral pores, his identification is undoubtedly correct. Probable sympatry, perhaps as widespread there as in the Tehuacan area, is suggested by the specimens of S. horridus available from Izucar de Matamoros (KU 43660), 4 mi E Matamoros (KU 38171), 6.5 mi SW Matamoros (KU 39631-4) and 1 mi SWTilapa (KU 38172). V ■ * ^ . f Although there is a paucity of records of occurrence of S. spinosus in northern Oaxaca, sufficient material is now available to demonstrate conclusively that its range in southern Puebla and northern Oaxaca broadly overlaps that of S. horridus, which accordingly is specifically distinct from S. spinosus. The difference in femoral pore number (>6 in the latter, <7 in the former) appears to be infallible, and the correlated absence of presence of contact of supraoculars with the median head scales is reliable to at least the 95% level. B. The Subspecies Groups (Exerges) of S. spinosus All members of the spinosus group from the state of Oaxaca, excepting S. horridus, S. melanorhinus and S. edwardtaylori belong to S. spinosus. S. horridus has fewer femoral pores (<7), S. melanorhinus has more numerous femoral pores (> 16, all in one row), and S. edwardtaylori has hypertrophied supraoculars, the rear ones of which contact both the median head scales and the superciliaries. Other trenchant differences exist among these four species. We have examined 139 specimens of S. spinosus from Oaxaca, as follows: vicinity of Cd. Oaxaca (UTA 8875-6; USNM 47218, 47346-8, 47534- 4; AMNH 18587-97, 18753-65, 18827-43); 1 1 mi E Oaxaca (UCM 16795-7); 13 mi E Oaxaca (UCM 8980-8, 49666-7); 25 mi E Oaxaca (UCM 8989-93); 25 mi S Oaxaca (UCM 8994-8); 37.7 km S Oaxaca (UTA 3389); Cerro San Felipe, nr Oaxaca (UCM 41130); Cerro San Pedro, nr Oaxaca (FMNH 1 10587); San Felipe del Agua, nr Oaxaca (UIMNH 60447-51); Tlacolula to San Pablo Mitla (AMNH 18804- 14); 3 mi W Mitla (KU 48998-45000, 40672- 3); 3.7 mi NE Mitla (UTA 4323); 0.5 mi NE Suchilquitango (32 km NW Oaxaca) ( UCM 16798); 0.5 mi S Guelatao (UCM 38364); El Tejocote (UTA 8602, 1 1804-5); Ejutla to Miahuatlan (AMNH 18598-9); Miahuatlan (AMNH 18721-5, 18822-6); 4 mi S Miahuatlan (UCM 45613-4); 6.6 mi S Miahuatlan Page 46 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 (UCM 48944); 6.8 mi S Miahuatlan (UCM 48945-7); 7.9 mi S Miahuatlan (UCM 48948); 15,7 mi S Miahuatlan (UCM 50724); 1 mi N Miahuatlan (UCM 22966-7); Cerro Yucuyacua, 8 km SSE Tlaxiaco (UCM 56476-84); 5.6 km WSW Tlaxiaco (UTA 2573, 2575-7, 3291). The one variable most nearly constant geographically in these specimens and in others reported in the literature is the number of supraoculars; four (on each side) in northern localities, and five in southern ones. Four supraoculars occur on each side in all three specimens from the northernmost reported locality of occurrence of S. spinosus in Oaxaca, El Tejocote. These conform in all respects with the features of the nominotypi- cal subspecies, to which they are referred. The single specimen from near Guelatao is also referred to the nominotypical subspecies. Larger series are needed from both localities to confirm these allocations, and additional material from northern Oaxaca in general is much needed to flesh out the distribution there of this species and subspecies. South of the localities represented by S. spinosus , 80% of the supraocular counts (sides separate, n = 130) are five or six, whereas north of Oaxaca (Smith, 1939) only 22% (n = 174) are more than four. One geographical irregularity evident in distribution of the occurrence of four supraoculars is in the Tlaxiaco (both UCM and UTA) series of 14 specimens, in which four supraoculars occur in 50% of the 28 sides. Excluding that series, 87% of the more southern S. spinosus have five or six supraoculars. We interpret the Tlaxiaco series as intergrades between S. s. spinosus , typically with four supraoculars, and the more southern subspecies, typically with five. Another sample of intergrades is available from near Mitla (3 mi W, and 3.7 mi NE; see list). In those six specimens four supraoculars occur on both sides of two, five in the other four. A third sample of intergrades is from 25 mi E Oaxaca; two of the five specimens have four supraoculars (both sides), the others five. The tendency in southern and central Oaxacan S. spinosus to have five or more supraoculars is accompanied by a high proportion (55% of 49 for which the character-state was recorded) of individuals with one or more supraoculars on one or both sides with a small lateral scale split off, giving rise to a lateral row of 1-5 small supraoculars. Such variants rarely occur in S. s. spinosus , and are not found in the northern Oaxacan examples here reported of that subspecies. In all Oaxacan specimens of S. spinosus, the supraoculars are usually all completely separated from the median head scales (80% of 138 sides), with exceptions widely scattered geographically, although nearly half of them ( 1 2 of 28) occur in the series of nine specimens in UCM from Tlaxiaco. Bulletin of the Maryland Herpetological Society Page 47 Volume 28 Number 2 June 1992 It was this series, and this variation, that led Boyer et al. (1983) to conclude that intergradation of S. spinosus and S. horridus was indicated, since contacts of supraoculars and median head scales are characteristic of S. horridus. As shown in the preceding account, however, that character is not infallibly diagnostic of S. horridus ; only the number of femoral pores is, in comparison with S. spinosus. The femoral pores of the Tlaxiaco series are typical of S. spinosus , 8- 1 1 (x = 9.7, n - 18). In addition, in a series of five from nearby, 5.6 km WSW Tlaxiaco (UTA 2573, 2575-7, 3291) not a single specimen has any supraocular in contact with the median head sc ales. The UCM series from Tlaxiaco appears to be anomalous in respect to its supraocular contacts. On the basis of number of supraoculars, the central and southern populations of S. spinosus are justifiably regarded as taxonomic ally distinct at the subspecific level, with intergradation evident both east and west of the central Oaxaca highlands. A marked geographic variation occurs within the population with more numerous supraoculars. Those in the vicinity of Cd. Oaxaca, according to Smith and Smith (1951) have more numerous dorsals (3 1 or more in 89% vs. 9%) and femoral pores ( 1 0 or more in 90% vs 1 3%) than the others; those two populations were designated S. s. caeruleopunctatus and S. s. apicalis. respectively. They constitute a 5-6 supraocular exerge distinct from the four supraocular exerge so far containing only S. s. spinosus. C. S. s. caeruleopunctatus Our data support the validity of two subspecies in the 5-supraocular exerge, although S. s. caeruleopunctatus appears to have an extremely small range in the vicinity of Cd. Oaxaca, and southward to 1 mi N Miahuatlan. Seventy-seven specimens we have examined clearly represent that subspe¬ cies: Oaxaca (49), San Felipe de Agua (5), Tlacolula to San Pablo Mitla (11), Ejutla to Miahuatlan (2), 1 mi N Miahuatlan (2), Cerro San Pedro (1), Cerro Felipe (1), 32 km NW Oaxaca (1), and 25 mi S Oaxaca (5). Only four (of 73) have 30 or fewer dorsals (28-37, x 32.7), and only nine femoral pore counts (in 149) are 9 or fewer (7-17, x 1 1.3). Several individuals have one or two extra, short rows of femoral pores (hence the record 17). D. S. s. apicalis S. s. apicalis has a range peripheral to that of the preceding subspecies, except to the north, where S. s. spinosus occurs. The inter¬ grades previously mentioned with-the latter subspecies involve only S. s. apicalis , as indicated by the frequent occurrence in them of only four supraoculars. Their dorsal and femoral pore counts, however, agree with Page 48 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 those of S. s. apicalis (as well as S. s. spinosus ), not with those of S. s. caenileopunctatus . Because at least a majority of the intergrades have five or more supraoculars, we allocate those specimens to S. s. apicalis. No intergradation of S. s. caendeopunctatus with S. s. spinosus is apparent in the material examined, and probably does not occur, contraiy to earlier as¬ sumption. Localities of record for S. s. apicalis are as follows (numbers are given of those examined in this study); Miahuatlan (10); 4 mi S, 6.6 mi S, 7.9 mi S, 15.7 mi S Miahuatlan (8); 8 mi S, 10 mi S Miahuatlan (Smith and Smith, 1951); San Pedro Quiechapa (op. cit.); 8 km SSB Tlaxiaco (9); 5.6 km WSW Tlaxiaco (5); 3 mi W Mitla (5); 3.7 mi NE Mitla (1); 25 mi E Oaxaca (5); 37.7 km S Oaxaca (1). The 43 specimens presently examined have 26-36 dorsals (x 29, n = 43, 7 (16%) >30), and 7-12 femoral pores (x 9, n = 86, 26 (30%) >9). The high frequency of femoral pore counts exceeding 9 is borderline for subspecific distinction, but the difference in dorsal scale count (5.5% with <31 dorsals in S. s. caendeopunctatus, 84% in S. s. apicalis) is not; respectively the femoral pore count is 6% <10 and 67% < 10). In the Miahuatlan area the transition from one subspecies to the other appears to be abrupt; at one mi N Miahuatlan, the two available specimens have 31 and 32 dorsals, and 10 or 11 femoral pores, and the two specimens from between Ejutla and Miahuatlan have 32 and 33 dorsals, 10-12 femoral pores. All specimens taken north of Miahuatlan along the highway to Cd. Oaxaca appear to represent S. s. caenileopunctatus , whereas those taken at Miahuatlan and southward represent S. s. apicalis. An exception is the specimen from 37.7 km S Oaxaca, which is clearly S. s. apicalis (28 dorsals, 8-8 femoral pores, 4-4.5 supraoculars. Presumably S. s. apicalis occurs westward towards Tlaxiaco, since we interpret the latter as intergrades with S. s. spinosus , but no specimens are yet available from that little-explored area. To the northeast, the subspecies occurs at San Pedro Quiechapa (Smith and Smith, 1951), and as intergrades with S. s. spinosus at 25 mi E Oaxaca and near Mitla (3 mi W, and 3.7 mi NE). Those eleven intergrades have 8 sets of 4 supraoculars, 14 of 5; only 2 femoral pore counts greater than 9; and only one dorsal scale count greater than 30. They clearly do not constitute intergrades with S. s. caenileopunctatus , of either S. s. spinosus or S. s. apicalis. Presumably the latter occurs in typical form in the area between San Pedro Quiechapa and Mitla, but no material from there is yet available except perhaps the specimen from 37.7 km S Oaxaca. Incontrovertible intergrades between S. s. apicalis and S. s. caemle- Bulletin of the Maryland Herpetological Society Page 49 Volume 28 Number 2 June 1992 opunctatus occur 1 i and 13 mi E Oaxaca. In 14 dorsal counts, 43% are <3 1 (26-35, x 30.9), and in 28 femoral pore counts, 50% are <10 (7-14, x 9.6). The intricate interdigitation of S, s. caeruleo p unc talus, S. s. apicalis , and their intergrades and intergrades of S. s. apicalis with S. spinas us east of Cd. Oaxaca (see Fig. 1) is no doubt correlated with an equally intricate interdigitation of habitats that merits study in detail. However, subspecific allocations based on series of one or two are inevitably suspect; series of six or more permit reasonably secure identifications. Key to the Subspecies of S. spinosus 1 . A. Supraoculars four on each side, seldom subdivided; dorsals seldom more than 30; femoral pores usually (83%) no more than 9 on each side, rarely more than 10 (3%) . . . spinosus B. Supraoculars 5-6 on each side, usually subdivided (Sceloporus spinosus exerge caeruleopunctatus) . . . 2 2. A. Dorsals usually no more than 30; femoral pores usually no more than 9 on each side . . . . . apicalis b. Dorsals usually 31 or more; femoral pores usually 10 or more, occasionally in 2 or more rows . . . . . .caeruleopunctatus Acknowledgments We are much indebted to the authorities in charge of the various collections from which material has been borrowed, including (in alphabeti¬ cal order of their collection acronyms) Drs. Charles W. Myers, Alan E. Leviton, Alan Resetar, John E. Simmons, Shi Kuei Wu, Christopher A. Phillips, Greg Schneider and Jonathan A. Campbell. We are also grateful for provisions of facilities for study at the University of Colorado, through the chairman of the Department of EPO Biology, Dr. William M. Lewis. Page 50 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 E Bulletin of the Maryland Herpetological Society Page 51 Volume 28 Number 2 June 1992 Literature Cited Boyer. Thomas H., Hobart M. Smith and Gustavo Casas Andreu. 1983. The taxonomic relationships of the Mexican lizard species Sceloporus horridus . Bull. Maryland Herp. Soc., 18: 189- 191 (1982). Cope, Edward Drinker. 1885. A contribution to the herpetology of Mexico. Proc. Am. Philos. Soc., 22: 379-404, fig. 9. Leviton, Alan E. 1980. Museum acronyms - second edition. Herp. Rev., 11:93-102. Smith, Hobart M. 1939. The Mexican and Central American lizards of the genus Sceloporus. Zool. Ser. Field Mus. Nat. Hist., 26: 1-397, fig, 1-59, pis. 1-31. Smith, Philip W. and Hobart M Smith. 1951. A new lizard ( Sceloporus ) from Oaxaca, Mexico. Proc. Biol. Soc. Washington, 64: 101-103. Department of Environmental Population and Organismic Biology, University of Colorado, Boulder, Colorado, 80309-0334 (HMS); and Department of Psychology, ibid., 80309-0345 (DC). Received: 26 September 1991 Accepted: 7 October 1991 Page 52 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 A REDESCRIPTION OF THE LARVAE OF RANA MONTEZUMAR (ANURA: RANIDAE) John K. Korky and Robert G. Webb Larvae of the Mexican frog Rana montezumae are redescribed. Most features do not differ appreciably from the description by Taylor (1942). The tooth-row formula of most larvae is 2(2)/3(l). Features of note in our sample of 30 tadpoles from near Ibarra, Guanajuato, include occurrence of a narrow P-1 medial gap (73%), a relatively short P-3 row about 63 percent the length of the P-2 row, an A-3 row never present, a wide A 2 medial gap (wider than either lateral segment), and occasional (23.3%) complete absence of the A- 2 row. Introduction Taylor (1942) first described the larvae of Rana montezumae based on two samples, one of 35 from the Distrito Federal containing both pre and post metamorphic individuals, the other of 9 tadpoles from the state of Mexico containing both “young and half grown” individuals. He described in detail a single larva of 112 mm total length from the larger sample; his illustration of this larva in lateral view ( 1 942:55, fig. 1) depicts well developed hindliinbs, but no exposed forelimbs, that mark stage 42 . Altig and Johnston (1986) listed this same larva at stage 36, which appears too early to judge from the hindlimb digits of Taylor’s figure. Taylor also illustrated the oral apparatus of this one larva (1942:55, fig. 5), and remarked about its papillae, musculature, and color pattern. The tooth row formula of 2(2) /3, discerned from Taylor's figure, agrees with that listed by Altig and Johnston for R. mon¬ tezumae tadpoles. Although Taylor included comments about individuals other than the illustrated larva, they are of a cursory nature, and statistical data were not employed. The purpose of this study is (1) to provide descriptive and statistical data on the morphology of 30 tadpoles of R. montezumae, and (2) to offer some characters that in combination will hopefully aid in identifying larvae of R. montezumae. Material^ and Method? A lot of 30 tadpoles (UTEP 7509, Laboratoiy for Environmental Biology, The University of Texas at El Paso) was seined by one of us (Webb) Bulletin of the Maryland Herpetological Society Page 53 Volume 28 Number 2 June 1992 and Peter Dalby, 20 July 1966, from a large shallow pond (maximal depth about 76 cm or 2.5 ft.) having abundant aquatic vegetation and aquatic insects. In addition to the ranid tadpoles, six salamander [Amby stoma) laivae (UTEP 7508) and one transformed ranid frog identified as Rana montezumae (Michigan State University Museum, Number 9587) were obtained at this pond site. The site of collection is on the property of Rancho La Puerta Guadalupe, about nine air kilometers southwest Ibarra, about 2560 m (8400 ft.), Guanajuato (near Jalisco border), Mexico. Larvae were staged according to Gosner (1960). Descriptive features follow Altig (1970) and Scott and Jennings (1985). Measurements of body length, tail length, tail musculature height, tail height, dorsal and ventral fin height, interocular and internareal distance, were made with Cenco calipers to the nearest mm, whereas those of “tooth” rows that include A-l length, left and right A-2 lengths, A-2 gaps, and lengths of P- 1 , P-2, P-3 and P- 1 gap, and mouth width, were made with a binocular dissecting microscope and ocular micrometer calibrated to the nearest 0. 1 mm. These morphometric characters were recorded for each of the 30 larva (missing or damaged parts in some individuals). Collective data for each character were subjected to univariate statistical analysis using an IBM personal computer and Axum (1989) statistical program, version 1.01. Descriptions of color pattern, thickness of the rectus abdominis muscle, body shape, lateral line system, and orbitonasal canal, follow Scott and Jennings (1985). Description Our 30 larvae vary from 32.0 to 87.0 mm in total length in developmental stages 27 to 41; the relationship of total length and stage of development are correlated in Table 1 ( a correlation coefficient between these two variables was determined asr = 0.95). Ascattergram (Fig. 1) shows a positive correlation between developmental stage and body length. The relative lengths of body and tail do not seem to vary with increasing size; tails are longer than bodies with the ratio tail/total length in 30 larvae varying from 0. 55 to 0. 63 (x , 0. 588) . The data for the 20 characters used in this study are indicated in Table 2. The body shape in dorsal view is bluntly triangular anteriorly with slight indentations posterior to the eyes that tend to separate the head from the abdomen at the anterior terminus of the dorsolateral muscles; this separation is obvious in lateral view, especially in the larger larvae. The proximal tail musculature is thick and robust. Color and pattern - Live tadpoles show considerable variation in colors and patterns within the same species due to ontogenetic and environmental variables. Hillis (1982) noted the effects of environmental factors on morphological features of both field-collected and laboratory- Bulletin of the Maryland Herpetological Society Page 54 Volume 28 Number 2 June 1992 Table 1. Relationship of size (total length) and Gosner stages of development in larvae of Rana montezumae; data are stage, sample size, mean, and range (mm). 27 2 34.5 32-37 28 2 39.5 35-44 29 1 45.0 45 30 31 4 49.5 48-53 32 1 47.0 47 33 2 55.0 54-56 34 3 60.6 54-67 35 2 62.5 61-64 36 2 69.0 67-71 37 4 80.5 76-87 38 2 73.0 73 39 1 82.0 82 40 3 81.3 79-84 41 1 84.0 84 Bulletin of the Maryland Herpetological Society Page 55 Volume 28 Number 2 June 1992 m O < h c/> ui S a. a -j 01 > 111 D H10N31 AQOe $ Page 56 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Table 2. Comparison of Means ± 1 SD and Ranges of 20 Morphological Characters of Field -collected Rana montezumae Larvae, N = 30, except as indicated. Character (in mm) (except #20) X ±1 SD (range) 1. Body length 25.30 ±5.62 (14.0-33.0) 2 Tail length 37.07 ±10.93 (18.0-57.0) a Total length 62.37 ±16.33 (32.0-87.0) 4 Tail musculature height 7.77 ±2.56 (3.0-12.00) a Tail height 14.03 ±3.60 (7.0-19.0) a Dorsal fin height 4.93 ±1.41 (3.0-9.0) 7. Ventral fin heigth 3.80 ±1.0 (2.0-6. 0) & Interocular distance 4.97 ±1.47 (3.0-9.0) 9. Intemareal distance 3.43 ±0.68 (2.0-5.0) 10. A- 1 tooth row length 3.41 ±0.82 (2. 1-5.0) 11. Left A-2 tooth row length 0.46 ±0.36 (0-1.3) 12. Right A-2 tooth row length 0.41 ±0.35 (0-1.3) 13. A-2 median gap width 1.34 ±1.0 (0-3.0) 14 A-2 gap ration (N=21) 0.37 ±0.19 (0.17-0.87) 15. P 1 tooth row length 2.96 ±0.73 (1. 8-4.2) ia P-2 tooth row length 2.99 ±0.73 (1.9-4. 1) 17. P-3 tooth row length (N=29) 1.89 ±0.61 (0.6-2. 7) la P-1 median gap width (N=22) 0.12 ±0.09 (0.1-0.5) 19. Mouth width 3.41 ±0.83 (2. 1-5.0) 20. Stage 34.30 ±4.13 (27-41) Bulletin of the Maryland Herpetological Society Page 57 Volume 28 Number 2 June 1992 reared larvae. Preserved tadpoles in formalin lose much of their color due to leaching, but retain melanin patterns in varying degrees. Scott and Jennings (1985) reported that the body wall of preserved tadpoles of both R . hlairi and R. pipiens becomes transparent, thus exposing the underlying melanin pattern of either the peritoneum, the rectus abdominus muscle, or the intestine. They used color and pattern to partly distinguish larvae of five species of New Mexican leopard frogs. Our preserved tadpoles of R . montezumae show variable retention of melanin depending on size. Small tadpoles tend to have clear, patternless tails, with reduced melanin in the body and tail musculature, and have indistinctly outlined body and tail myomeres. Features of color and pattern are best developed in the large tadpoles that have the body dorsum and sides dark gray with distinctly outlined dorsolateral myomeres; the venters are generally clear or light gray with the intestines visible. The pale yellow tail musculature has distinct, mostly patternless, myomeres anteriorly, but a fine mottling of coalesced dots and flecks posteriorly. The fins, clear anteriorly, have spots and flecks, and in some larvae a guanic reticulation, posteriorly with the pigmentation often most intense in the lower fin. Lateral line organs and orbitonasal canal - These features were observed at 7-X0x magnification under water with a binocular dissecting microscope. The orbitonasal canal is visible only in large tadpoles (stage 38- 41). The lateral line system seems to be mostly absent even in the large tadpoles; only parts of the dorsal and lateral branches are evident as discontinuous stretches of pellucid dots. Oral disc - The sides of the oral disc are emarginate. The configura¬ tion of oral papillae and the homy beak do not differ appreciably from the description by Taylor (1942); the papillae, pigmented or not, are small and sparse inside the comers of the mouth. The tooth-row formula applicable to most larvae is 2(2)/3(l). The length of the A- 1 tooth row is about the same length as the width of the mouth. The A- 2 tooth row is variable. Both right and left segments are separated by a wide and variable median gap (wider than either lateral segment) in 2 1 of 30 larvae (70%) for which the mean A- 2 gap ratio is 0.370 (0.17-0.87). In two larvae (6.6%) only the right or left segment of the A-2 row is present (absent on other side), whereas seven larvae (23.3%) lack both right and left segments (A-2 row absent). This variation in the A-2 row does not seem to be correlated with the stage of development (Table 3). None of our larvae has a third upper tooth row IA¬ SI . A narrow medial gap (only 0. 1 to 0.5 mm) in P- 1 in 22 of 30 larvae (73%) is not indicated by Taylor (1942) or Altig and Johnston (1986). The third lower tooth row (P-3) is noticeably shorter than P-1 or P-2 (about same length), as depicted by Taylor (1942; 55, fig. 5). One larva of 30 lacks a P-3 Page 58 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Table 3. Number of tooth rows in upper labium (A- 1 , A-2) showing variation in A- 2 (1.5, one segment absent: 2+ , both segments present; 2-, both segments absent). Numbers of tooth rows Stage N 1 1.5 2(+) 2() 27 2 28 2 29 1 30 31 4 32 1 33 2 34 3 35 2 36 2 37 4 38 2 39 1 40 3 41 1 2 2 1 1 right 1 2 1 4 1 2 3 2 2 4 2 1 3 1 1 left 2 1 3 2 3 1 1 3 1 1 1 1 2 1 1 Bulletin of the Maryland Herpetological Society Page 59 Volume 28 Num ber 2 June 1992 Fig. 2. Oral disc of larva of Raria montezamae , Gosner stage 34, 54 mm total length (UTEP 7509) Page 60 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 row. The mean length of the P-3 row in 29 larvae is 1.89 (0.6-2. 7) mm compared to 2.99 (1.9-4. 1) mm for the P-2 row in 30 larvae, or the mean length of row P-3 is about 63 percent the length of P-2. The oral disc of R. montezumae is illustrated here in Figure 2. Comparisons Detailed descriptions of anuran larvae have purpose when they can be ultimately used for comparison by future investigators. Suitably detailed descriptions are not available for taxa of frogs of the Rcma pipiens complex from areas of sympatry or near allopatry with R. montezumae. Despite the geographical disparity of the larval samples, our data are in general agreement with that of Taylor (1942) and suggest the following combination of characteristics of R. montezumae as possibly useful in comparing larvae of frogs of the R. pipiens complex (based on comparative data in Scott and Jennings, 1985); (1) the relatively dark-blotched pigmentation posteriorly on tail, (2) an A-3 tooth row never present, (3) a wide A-2 gap (wider than either lateral tooth-row segment), (4) occasional absence of the A-2 row, (5) the relatively short length of row P-3. When compared to larvae of the five species of the R. pipiens complex from New Mexico (Scott and Jennings, 1985), the features of R. montezumae most closely match those of R. chiricahuensis, and thus support inclusion of these two species in the R. montezumae species group (Hillis et al., 1983; 1988). Bulletin of the Maryland Herpetological Society Page 61 Volume 28 Number 2 June 1992 Fig. 3. Representative larvae showing features of color and pattern best devel¬ oped in larger tadpoles (UTEP 7509). Scale equals 40 mm. Page 62 Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 Acknowledgments We are indebted to: Mr. John Smalley of Montclair State College for his assistance with statistics and graphics; Mr. David Fogg for his assistance with photography; and the Montclair State College Separately Budgeted Research Program for support. Literature Cited Altig, R. 1970. A key to the tadpoles of the United States and Canada. Herpetologica, 26; 180-207. Altig, R. and G. F. Johnston. 1 986. Maj or characteristics of free-living anuran tadpoles. Smith¬ sonian Herpetol. Inform. Service, (67): 1-75. Axum. 1989. Gosner, K. L. 1960. Hillis, D. M. 1982. Hillis, D. M. 1988. August, First edition,version 1.01, Trimetrix, Inc., 444 N.E. Ravenna Blvd., suite 210, Seattle, WA 98115. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16: 183-190. Morphological differentiation and adaptation of the larvae of Rana berlandieri and Rana sphenocephala {Rana pipiens complex) in sympatry. Copeia, 1982 (1): 168-174. Systematics of the Rana pipiens complex: puzzle and para¬ digm. Ann. Rev. Ecol. Syst., 19: 39-63. Hillis, D. M., J. S. Frost, and D. A. Wright. 1983. Phylogeny and biogeography of the Rana pipiens complex: a biochemical evaluation. Syst. Zool., 32: 132-143. Scott, Jr., N. J. and R. D. Jennings. 1985. The tadpoles of five species of New Mexican leopard frogs. Occas. Papers Mus. Southwestern Biol., Univ. New Mexico, (3): 1-21. Bulletin of the Maryland Herpetological Society Page 63 Volume 28 Number 2 June 1992 Taylor, E. H. 1942. Tadpoles of Mexican anura. UK Sci. Bull. 28: 37-55. Biology Department Montclair State College , Upper Montclair , New Jersey 07043 and Department of Biological Sciences, University of Texas at El Paso, El Paso, Texas 79968-0519. Received: 17 October 1991 Accepted: 28 October 1991 Page 64 i Bulletin of the Maryland Herpetological Society Volume 28 Number 2 June 1992 AN ALBINO THAMNOPHIS SIRTAUS FROM FAIRFIELD COUNTY, SOUTH CAROLINA Instances of albinism in Thamnophis sirtalis have been reported numerous times in the literature. Dyrkacz (1981) cites at least seven examples of albinism in this species in the SSAR Herpetological Circular No. 1 1 : Recent Instances of Albinism in North American A mphibians and Reptiles. On September 1, 1991 an albino T. sirtalis was collected in a residential area of Fairfield County, South Carolina by a local individual and presented to the Riverbanks Zoological Park, Columbia, South Carolina for identification. The specimen, an adult female, is ca. 66 cm in length and exhibits similar coloration as a juvenile collected by Cope and Kashner near Dansville, Montour County, PA during 1974 and reported by Groves [( 1976) Bull. Maryland Herp. Soc. 24: 21-22]. This individual is currently being maintained in the collection of the Riverbanks Zoological Park, Columbia, South Carolina. Upon death, it will be deposited in the Vertebrate Collection of the South Carolina State Museum, Columbia, South Carolina. —Chuck Smith Department of Herpetology, Riverbanks Zoological Park, Columbia, South Carolina —Kyle SchuetU Department of Herpetology, Riverbanks Zoological Park, Columbia, South Carolina Received: 18 September 1991 Accepted: 18 September 1991 Bulletin of the Maryland Herpetological Society Page 65 Volume 28 Number 2 June 1992 Bulletin of the Maryland Herpetological Society Page 66 Volume 28 Number 2 June 1992 ‘MOVES REPTILE & AMPHIBIAN SHOW and SALE to be held Saturday, November 14 & Sunday, November 15 9 a.m. to 5 p.m. 1 0 a.m. to 5 p.m. in Bing Crosby Hall at the Del Mar Fairgrounds, Del Mar, California (1-5 to Via de la Valle exit west, Left on Jimmy Durante Blvd to main fairgrounds entrance.) 12 years to adult - $3.00 NOTICE: Due to fairground regulations, only animals belonging to participants or those purchased at the show will be allowed on the grounds, with the exception of guide dogs. For further information, call (619) 689-2582 Live Amphibian & Reptile Exhibits Educational Displays Animals and Animal-care Products, Books, Posters, T-shirts, Art, Jewelry, and much more will be offered for sale by a variety of vendors Bulletin of the Maryland Herpetological Society Page 67 Volume 28 Number 2 June 1992 9&PES Venomous Reptiles of North America New Titles Carl H. Ernst H Natural History H Herpetology An up-to-date and definitive natural history of all venomous reptiles north of Mexico, Venom¬ ous Reptiles of North America presents these shy, beautiful, but often maligned creatures as an integral pan of the ecology of our continent. Invaluable to herpetologists, wildlife biolo¬ gists, naturalists, and fans of the outdoors, this book is the first comprehensive survey of these species in more than thirty years. It describes the biology and natural history of the twenty venomous snakes and one lizard found in the United States and Canada. Introductory discus¬ sions cover external morphology, venomology, and conservation. An in-depth identification key helps reduce the difficulties in correctly identify¬ ing venomous reptiles. Line drawings, photo¬ graphs, color illustrations, and maps of species distributions further aid the reader. Detailed species accounts include a description, karyo¬ type, fossil record, distribution, geographic variation, notes about confusing species, habitat, behavior, reproduction, growth and longevity, food habits, venom and bites, predators and defense, and conservation status. Because habitat destruction by humans is the most common cause of the decline of these animals, Ernst focuses on their conservation and protection. He also presents a pronunciation glossary of scientific names and definitions, an extensive bibliography directing wildlife biolo¬ gists and naturalists to additional references of interest, and a taxonomic and subject index. 9 Smithsonian Institution Press Car! H. Ernst is professor of biology at George Mason University, Fairfax, Va. He has written four books, including the award-winning Turtles of the World (Smithsonian Institution Press, 1989). 55 color, 33 b&w Ulus. 7 x JO 216 pp. LC 91-3535 Cloth: ISBN 1 -56098- 1 1 4-8H $3 5. 00 July 1992 Rights: World Spring 1992 Page 68 Bulletin of the Maryland Herpetological Society Society Publication Back issues of the Bulletin of the Maryland Herpetological Society, where available, may be obtained by writing the Executive Editor. Alist of available issues will be sent upon request. Individual numbers in stock are $2.00 each, unless otherwise noted. The Society also publishes a Newsletter on a somewhat irregular basis. These are distributed to the membership free of charge. Also published are Maryland Herpetofauna Leaflets and these are available at $.2 5 /page. Information for Authors All correspondence should be addressed to the Executive Editor. Manuscripts being submitted for publication should be typewritten (double spaced) on good quality 8 1/2 by 11 inch paper with adequate margins. Submit original and first carbon, retaining the second carbon. If entered on a word processor, also submit diskette and note word processor and operating system used. Indicate where illustrations or photographs are to appear in text. Cite all literature used at end in alphabetical order by author. Major papers are those over 5 pages (double spaced, elite type) and must include an abstract. The authors name should be centered under the title, and the address is to follow the Literature Cited. Minor papers are those papers with fewer than 5 pages. Author’s name is to be placed at end of paper (see recent issue). For additional information see Style Manual for Biological Journals (1964), American Institute of Biological Sciences, 3900 Wisconsin Avenue, N.W., Washington, D.C. 20016. Reprints are available at $.03 a page and should be ordered when manuscripts are submitted or when proofs are returned. Minimum order is 100 reprints. Either edited manuscript or proof will be returned to author for approval or correction. The author will be responsible for all corrections to proof, and must return proof preferably within 7 days. The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland , Inc. 2643 North Charles Street Baltimore, Maryland 21218 US ISSN: 0025-4231 EPT BULLETIN OF THE ^Rarylanb f)erpetological ©oriety DEPARTMENT OF HERPETOLOGY THE NATURAL HISTORY SOCIETY OF MARYLAND, INC. cM\ fl m o 2 \m ?ARi MDHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League 30 SEPTEMBER 1992 VOLUME 28 NUMBER 3 BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY Volume 28 Number 3 CONTENTS Swptember 1992 "Shouldering" in Prairie Rattlesnakes: A New Hypothesis David Chiszar, Jason Perelman, Hobart M. Smith and David Duvall . 69 Rattle Length in Crotalus horridus atricaudatus Charles Smith . 77 New Hatchling Mud Snakes, Farancia abacura reinwardtii Schlegel Harold A. Dundee . 78 A New Chormospecies of Snake (Pseudoleptodeira) from Mexico Aurelio Ramirez Bautista and Hobart Smith . . 83 Additional Rainbow Snakes, Farancia erytrogrammcc from Charles County, Maryland Robert Miller . 99 ! The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc. 2643 North Charles Street Baltimore, Maryland 21218 BULLETIN OF THE Volume 28 Number 3 September 1 992 The Maryland Herpetoiogical Society Department of Herpetology, Natural History Society of Maryland, Inc. Executive Editor . Herbert S. Harris, Jr. Steering Committee Frank Groves Jerry D. Hardy, Jr. Herbert S. Harris, Jr. Library of Congress Catalog Card Number: 76-93458 Membership Rates Membership in the Maryland Herpetoiogical Society is $16.00 per year. Foreign $20.00/year. Make ah checks payable to the Natural History Society of Maryland, Inc. Meetings The third Wednesday of each month, 8:15 p.m. at the Natural History Society of Maryland (except May — August, third Saturday of each month, 8:00 a.m.). The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 p.m. Volume 28 Number 3 September 1992 “SHOULDERING” IN PRAIRIE RATTLESNAKES: A NEW HYPOTHESIS1 David Chiszar, Jason Perelman, Hobart M. Smith and David Duvall Prairie rattlesnakes were tested with kingsnake chemical cues in either a pen or an artificial burrow. The dependent variable was frequency of body-bridging responses. More responses occurred to these chemical cues in the burrow situation than in the pen. Further, body bridging in the burrow can (1) give the rattlesnake firm purchase against the walls, and (2) seal the passage ways. Both factors could discourage predatory attempts by kingsnakes. Ac¬ cordingly, we hypothesize that body bridging might be adaptively associated with crevice and burrow situations, and this response might be better studied in these situations than in open arenas. Crotaline snakes exhibit a specialized reaction to kingsnake preda¬ tors (Cowles, 1938; Klauber, 1927), triggered by chemical cues associated with the integuments of these ophiophagous snakes (Bogert, 1941; Weldon & Burghardt, 1979). This behavior has received several names (e.g., shouldering, kingsnake defense posture, body bridging, inverted U), but all investigators agree in describing its essential features: “A body bridge starts from a normal laterally projecting loop of a portion of the trunk of the snake in resting positions and rises from the substrate. As the body bridge rises, the dorsoventral plane of the trunk region involved is tipped or tilted medially, thus presenting the ventral surface or belly laterally to the outside of the body bridge” (Carpenter & Gillingham, 1975, p. 295; see this article for many additional details and for excellent drawings). Other behaviors occur along with body bridging (e.g., inflation of the trunk, jerking move¬ ments of the trunk and of the raised bridge), apparently augmenting the display (Carpenter & Gillingham, 1975; Klauber, 1956). 1The behavior here under study has been called body bridging for many years, and we use this term in the text. The word “shouldering" is used in the title, however, to recognize our debt to Dr. Charles M. Bogert, with whom we have corresponded during the conception and execution of this study. Dr. Bogert’s (1941) contribution to this line of research is well known; indeed, it is a classic paper in herpetological ethology. Dr. Bogert has long referred to our target behavior as “shouldering", and we take pleasure in acknowledging our interaction with him by using his term in our title. Our text will make clear, however, that we are making no attempt to supplant the nomenclaturally- stable term body bridging. Bulletin of the Maryland Herpetological Society Page 69 Volume 28 Number 3 September 1992 Although there is no doubt about the reality of body bridging, the behavior is quite variable in that some individuals exhibit it readily while others do not; also, snakes that perform the behavior during initial stimu¬ lation frequently quit responding after a few trials (Bogert, 1941; Gutzke, 1991). As far as we are aware, no one has explained this variability or has developed a paradigm that eliminates it (although Gutzke, 1991, hypothe¬ sized that size of the crotaline is inversely correlated with frequency of body bridging to a kingsnake stimulus). This could mean that all studies of body bridging have used sub optima! setting conditions and that alternative con¬ ditions could give rise to less variability in the behavior. A characteristic common to all studies of body bridging has been that crotaline snakes were tested in arenas with little or no cover available. This fact became salient to the authors when we were collecting prairie rattle¬ snakes (Crotalus viridis) along sandstone outcrops near Agate, CO. One snake, a large female, managed to get half of her body into a crevice before we got our Pilstrom tongs on her. Although we captured this animal, approximately 20 minutes were required to dig her out. During the digging we saw that the snake’s body was bridged at several points and these bridges were pressed firmly against walls of the sandstone tunnel. Such appositions made the animal veiy difficult to remove, and at particularly narrow points in the tunnel the passage was completely obstructed by them. These observations led us to hypothesize, perhaps iconoclastically, that body bridging might be a behavior of greater utility to crotaline snakes in burrows or crevices than to the same snakes above ground. Indeed, it has never been abundantly clear that body bridging by a crotaline in an arena situation could actually deter a large kingsnake bent on eating the crotaline (see Klauber, 1956, for a different opinion; and see Carpenter & Gillingham, 1975, for an alternative hypothesis). On the other hand, it was our impression during the Agate collecting trip that a kingsnake would not be able to enter the rattlesnake’s lair because it was effectively sealed by the bridges. Furthermore, a kingsnake with a mouth grip on the rattlesnake’s midsection or rear would not be able to pull the firmly anchored prey out of the burrow. To test these ideas we constructed an artificial burrow and compared the elicitability of body bridging in rattlesnakes confined to the burrow with the elicitability of this behavior in snakes living in a pen that was comparable to arenas used in previous studies. If body bridging is a response that is especially effective in burrows, then we ought to see this behavior more frequently in rattlesnakes tested in the burrow than in snakes tested in the pen. To standardize the test, we use kingsnake chemical cues (presented on rods and cotton-tipped applicators) as stimuli rather than kingsnakes themselves. Whereas it would be difficult (perhaps impossible) to control the Page 70 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 behavior of a live kingsnake in the pen or the burrow, we could control precisely the movement and position of the rods and applicators. Methods Eight C. viridis (adult, long-term captives) were observed. Four were confined, individually, in the burrow (Fig. 1), and four were confined, individually, in a plywood pen (60 x 66 x 81 cm). A rattlesnake was placed into its test situation two days prior to the first observation. On day three, five tests were administered: (1) the tip of a plastic rod that had been rubbed along the back of a kingsnake was brought with 2 cm of the rattlesnake’s face, (2) a cotton- tipped applicator that had been rubbed along the back of a kingsnake was brought within 2 cm of the rattlesnake’s face, (3) the tip of a control plastic rod that had been dipped in tap water was brought within 2 cm of the rattlesnake’s face, (4) a cotton-tipped applicator that had been dipped in tap water was brought within 2 cm of the rattlesnake’s face, and (5) no disturbance. Tests were administered in random order with 30 min separating successive trials. Each test was 60 sec long, with the rod tip or applicator being moved as necessary to keep it within 2 cm of the rattlesnake’s face. This entire five-trial procedure was replicated five times at 24-hr. inter¬ vals. Hence, each snake had a total of 25 tests, five with each stimulus. The dependent variable was presence versus absence of at least one body¬ bridging response during each test. Two kingsnakes (one Lampropeltis getula splendida and one L. g. califomiae) were used as chemical cue donors. They were maintained in separate cages (62 x 32 x 32 cm), and neither had ever been brought into contact with the C. viridis used in this study. The kingsnakes were used equally often as chemical donors for each of the rattlesnakes. Chemical cues from the two kingsnakes were equally effective in eliciting body bridging responses (F < 1), so data were pooled across this factor and it will not be discussed further. When a snake was removed from the pen or the burrow, the apparatus was cleaned with water and toweled diy. A new snake was installed, and another two-day acclimation period passed before observa¬ tions commenced. The laboratory was maintained at 26°C by electric heaters and the photoperiod (0700-1900) was automatically controlled. All snakes normally lived individually in glass terraria (62 x 32 x 32 cm) containing paper floor covers and a stainless steel vessel filled with water. Feeding occurred once per fortnight (one mouse, Mas musculus , 20 g) . Snakes were fed prior to being Bulletin of the Maryland Herpetologicai Society Page 71 Volume 28 Number 3 September 1992 placed into the experimental apparatus and were not fed while they were in it. Results Body bridging was never seen during trials involving no disturbance, and this behavior was rarely seen with control applicators or with the control rod (Table 1). On the other hand, body bridging was frequently observed when kingsnake chemical cues were introduced (Table 1). Data for each snake were summarized by calculating percent of trials for each stimulus that contained a body-bridging response. Each of these five percents were then averaged over snakes in the burrow condition and in the pen condition. Analysis of variance then treated conditions (pen versus burrow) as a between-subj ects factor and stimuli as a repeated-measures factor. Be¬ cause percentages are frequently distributed in a skewed manner, we applied the ANOVA not only to the percents but also to arcsins and probits, commonly used transformations (Winer, 1971). All the ANOVAs revealed similar patterns of effects, and we here report the outcome of the probit analysis. Not surprisingly, the effect of stimuli was significant (F = 6.66, df = 4,24, P < 0.01). Post hoc contrasts revealed that kingsnake chemical cues on rods and applicators were equally likely to generate body bridging, and both types of tests had significantly more bridging responses than did the control or undisturbed trials (which did not differ among themselves). Rattlesnakes in the burrow environment emitted more body bridging responses than did snakes in the pen (main effect of burrow vs pen: F = 3.95, df =1,6, 0.10 > P > .05. More important, since this difference between burrow and pen was restricted to rods and applicators containing kingsnake chemical cues, the ANOVA produced a significant interaction between burrow vs pen and stimuli (F = 3.56, df = 4,24, P < 0.05). Discussion These data support the hypothesis that body bridging by prairie rattlesnakes is more likely to be seen in burrows than in open areas such as our pen. It is noteworthy that several of the bridging responses seen in the pen involved the rattlesnake looping its body against a wall, as if the snake was attempting to gain purchase. This was veiy similar to the body bridges seen in the burrow. Most important, body bridges in the burrow had the effect of pushing the snakes’ bodies against several walls simultaneously, and this would make the snakes difficult to pull out of the burrow. The Page 72 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 T3 V •e 3 g § CO o o •3 © d a D U O fc TJ Vh d T3 fi a p-i o 8 * i % g s fi3 o a O a o d o d m V *8 CO > §1 g CO a C 2 O o co 3 8-“ «M a o d d 5 o * S *3 * if u •d js al . 3 S b &£ | > §‘■3 a -a <2 •* p Ml III 4) al £.8 25-0 S> -■g § si: 13 ’ I’i® Q. fl d _SC|i ■§ - § al® a»ilfl 111!-.! ^®||l ** oi *J a iiliil 1123;! s=2;2i fastis B o *4? *3 o 4J HslISo s Jj ° & 0*3!^ 3S a 3 Slsii M«| ^ 2 o 8 75 9 a" si! 3 § “ g kd S °3^ 5 2" > k5 S’; *J rt -3 I S2al I sti lili 1*1 1 *5 'o > - s < g- a a a <; a 5 > o z § § S 5 g S a s 52 w 3 on 52 u- w C 3 w**2 3 g g § _< ^ § § A -Q ■Sf 11 Ed S..IS - Wj gss l! <3 < Cd z s ° hS o a a. < Page 104 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 Announcing: Biology of the Pitvipers Edited by Jonathan A. Campbell & Edmund D. Brodie, Jr. ★ 467 pages containing thirty contributions by 5 1 authors. ★ Treats systemancs. evolution, technique, natural history, behavior, snake-bite, maintenance and conservation. ★ 122 color photos, 100 black & white photos. ★ Case bound, with dust jacket. ★ Printed on acid-free paper. ★ Price: US $75.00 (includes shipping) ★ ISBN 0 - 9630537-0-1 "This imposing collection of herpeiological contnbuuons provides an outstanding overview of the most highly specialized of all the snakes." Roger Conans - Ad). Professor, The University of New Mexico “From a clinical standpoint . . . Biology of the Pitvipers is a timely addition to the American snakebite literature The sequelae for pirviper poisoning in the United States by RC Dan et al fills an important gap in the literature. It documents for the first time the long range effects of pitviper bites in a careful, systematic and scientific fashion. The entertaining and readable style in which this account is written is an added bonus. (David Hardy's) comprehensive evaluation of the Extractor® suction device and Stun Gun^ electroshock are the most in- depth and thorough review s to date of these controversial maneuvers." George Wda M.D.. DTM&H - Chief, Department of Medicine. AFRIMS "For the first time, the many studies of the spellbinding (pitvipers] are being pulled together into a single volume... The book . . . presents a sweeping view of the pn s iper s evolution, ns bizarre anatomy . ns alternately violent and gallant mating behavior, the lethal brew ol toxins and degradaliie enzymes found in ns venom, its singular approach to eating and other solutions the creature has devised to the problem of being a snake " The New York Tunes “This volume is a good investment lor serious students of viper evolution and ecology The editors have brought together a number of timely . solid contnbuuons that succeed admirably in providing both a good introduction summanes of previous work, and new data and ideas. Recent years has e brought us a number of other sympo¬ sium volumes, but few have achieved this general level of scholarship " Darrel R. Frost Ass'L Curator of Herpetology. American Museum of Natural History The cover illustration of a Cane brake Rattlesnake. Crotalus horndus. by William B Montgomery . was commissioned especially tor Biology of the Pitvipers It is a limited edition etching, hand printed on acid-fret archival paper, signed and numbered by the artist Each impression is individually hand colored by the arust using light-fast warercolors The 16" x 22 pnnts will be sold at a special discount pnee of $73.00 until August 15. 1992 (regular pnee SIOO OOi 13 • l S V vl'honc (9031 593-717(1 .FAX (9I»3> 597-5131 Cut here to mail or FAX ! j copy! s i of Biology of fhe Pilvipers for US $75.00 ppd i | Canebrake Rattlesnake pnnt(s) for US $75.00 ppd Book rate/surface postage paid worldwide Texas residents please add t sales lax FREE , Street or P O Bix □ Check or money order enclosed (US dollars only, pleasei payable to SELVA □ Viva □MasterCard Exp. dale _ Your signature auiltoni . Sel«a to teba sour cm charee wdl he added ir Zip or Postal Code Phone number/FAX Purchase orders and wholesale or dealer inquiries welcome. □ Please add me to your mailing list □ Send me information on uie cover illusirauon SELVA PO Box 5213 Tvler. Texas USA 75712-5213 Phone (903) 593-7170 F\\l(nRttRDLR:m.tL597-5l3l * FAX TOFU ORDER! (90.1) 5V7o/.?/ ★ ?★ * Bulletin of the Maryland Herpetological Society Page 105 Volume 28 Number 3 September 1992 o^lws & ‘Notes HERPETOLOGY Contemporary Research on the Biology of Amphibians and Reptiles HERPETOLOGY is ai once an old and yet also a newly revitalized field of research. It was founded in the 18th and 19th rwnnri« on taxonomic, acaasnicai, and distributional studies, hnl rlnring tbp rynfnry ffi-rvfo jr> biology anri crynfy frngiTV^itrod thr «tmrly nf gray&iiKirjris and reptiles into narrower and more disconnected disciplines. The recent renaissance of holistic approaches has helped to transcend these disciplinary boundaries, enabling herpetologists to become active p@tkipants and even leaders in this rebirth of the integrated study of animals, as the authors in this new volume demonstrate. The “New Herpetology” was formalized in 1989 at the First World Congress of Herpetology, held in the United Kingdom, which brought together more than 1000 specialists on amphibians and reptiles from some 60 countries. Ecologists, anatomists, conservationists, geneticists, physiologists, paleontologists, and others who span the spectrum of biology met to share ideas and information. There was a general acknowledgment of the value of broadly integrated approaches to the study off a groep of organisms, and the recognition that modem herpetology represents a vigorous and cohesive discipline of enduring significance. The main feature of the Congress was a senes of plenary (ceases chosen to emphasize the range of current berpeiological studies and to highlight those topics in which research on amphibians and reptiles have made major contributions to biology. This book contains the revised and updated versions of those lectures by authors who are among the world’ s leaders in their respective fields. Together, these essays demonstrate the breadth of modem herpetology and its continuing vitality as a discipline. The volume also contains a detailed. illustrated summary of the meeting, with a list of ail participants in symposia, workshops, roundtables, poster sessions, and other events. Included is a complete list of delegates and their addresses. This volume represents the summary of record for the Fust World Congress of Herpetology and a synopsis of current research in our discipline. It is an excellent introduction to modem herpetology for students and others interested in the biology of amphibians and reptiles. Specifications: 225 pages, format 8 */i by 1 1 inches (22 by 28 cm), 20 photographs, numerous tables and graphs, dotbbound in library-grade buckram. To be published December 1992. Table of Contents Carl Cans (USA): “Tie Status of Herpetology” ILYA S. Darevsky (Russia): “Evolution and Ecology of Parthenogenesis in Reptiles” LINDA MAXSON (USA): ‘Tempo and Pattern in Anuran Speciadon and Phylogeny: An Albumin Perspective” Russell A. MITTERMEIER (USA) AND OTHERS: “Conservation of Amphibians and Reptiles” TIM Haluday (UK): “Sexual Selection in Amphibians and Reptiles: Theoretical Issues and New Directions” ARMAND DE RICQLEs (France): “Paleoherpetology” S. Donald Bradshaw (Australia): “Eccphysioiogy of Desert Reptiles” ERIC R. PIANKA (USA): ‘The State of the Art in Community Ecology” David B. wake (USA): “An Integrated Approach to Evolutionary Studies of Salamanders” SUMMARY OF THE FIRST WORLD CONGRESS OF HERPETOLOGY LIST OF CONGRESS DELEGATES WITH ADDRESSES Prices and Ordering The pre-publication price (before 1 5 November 1992) to SSAR members sad to Delegates of the First Congress is $20. The price to all other persons and to institutions is $28. (Packing and shipping per volume: USA add $2, ocher countries add $4). Numbers of copies _ at $ _ per copy, plus $ _ total packing and shipping: total amount enclosed: $ Send orders to: Dr. Robert D. Aidridge. SSAR Publications Secretary, Department of Biology, St Louis University, St Louis, Missouri 63103, USA. (Telephone 3 14-658-3916; fax 3 14-658-3 117). Overseas customers must pay in US A funds using a draft drawn on American banks (include an additional amount to cover bank conversion charges) or by International Money Older. MasterCard or Visa are accepted (provide account number and expiration date); a 5% bank charge will be added to your accounL A complete list of SSAR publications and applications for Society membership are available on request from Dr. Aldridge. Page 106 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 9lffWS dr 9{OTES Second World Congress of Herpetology 29 December 1993 - 6 January 1994 For information please contact: Michael J. Tyler, Department of Zoology, University of Adelaide, Box 498, GPO Adelaide S.A. 5001, Australia. FAX: 61-8-223-5817 FOR SALE Herpetological Reprints . $0.25 ea. Call Joe Lewandowski at (301) 358-7834 RELEASE FOR IMMEDIATE PUBLICATION Upstate Herpetological Association is now accepting applications for a grant in the amount of $400.00 to be awarded at our November meeting. Qualified applicants will submit a proposal detailing research or study of reptiles or amphibians on or before September 15th to Upstate Herpetological Society, HCR68 Box 30B Springfield Center, NY 13468. New or ongoing projects will be considered equally, however, preference may be given to those projects involving species native to New York. Successful recipients will be required to submit a project summary for publication in the Herp Beat newsletter. Bulletin of the Maryland Herpetological Society Page 107 Volume 28 Number 3 September 1992 The International Herpetological Symposium, Inc. Announces the 17 th Annual Symposium to be held in Miami Beach, Florida June 17th through June 20th . CALL FOR PAPERS The IHS presents papers on herpetoculture, natural history, veterinary medicine and other topics related to herpetology. Individuals interested in presenting a paper at this meeting should write to Richard Ross,MD, Institute for Herpetological Research, PO Box 2227, Stanford, CA 94309. The International Herpetological Symposium, Inc. Announces the 17th Annual Symposium to be held in Miami Beach, Florida June 17th through June 20th. CALL FOR PAPERS The IHS presents papers on herpetoculture, natural history, veterinary medicine and other topics related to herpetology. Individuals interested in presenting a paper at this meeting should write to Richard Ross,MD, Institute for Herpetological Research, PO Box 2227, Stanford, CA 94309. The International Herpetological Symposium, Inc. Announces the 17th Annual Symposium to be held in Miami Beach, Florida June 17th through June 20th. CALL FOR PAPERS The IHS presents papers on herpetoculture, natural history, veterinary medicine and other topics related to herpetology. Individuals interested in presenting a paper at this usee ting should write to: Richard Ross,MD, Institute for Herpetological Research, PO Box 2227, Stanford, CA 94309. Page 108 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 O^E/WS &9&PES EXPEDITION TO TOBAGO, WEST INDIES January 4 to January 16, 1993 HERPETOLOGY ENTOMOLOGY MARINE BIOLOGY BOTANY This is part of an on-going survey of the flora and fauna of Tobago. This trip will emphasize photographing the Tobago reptiles, and collecting samples of a number of little-known plant and animal groups. Participants will be housed in a 10-bed, 4-bedroom beach house (2+2+2+4) which has an adequate kitchen (we share the task of cooking) and hot showers. The cost, which is $40 U.S. per day, includes all meals, housing, and shared rental cars (we usually rent three cars for a group of ten people). Scuba diving is available at additional cost Group air fares may be available out of the Baltimore-Washington area. For further information contact: Jerry D. Hardy, Jr. 22 Wade Avenue Baltimore, Md., 21228 Phone 301-747-7724 Bulletin of the Maryland Herpetological Society Page 109 Volume 28 Number 3 September 1992 9\@WS df 9iOT£S STEVEN SIMPSON NATURAL HISTORY BOOKS PO BOX 853 • BRIGHTON • BN1 5DY England Tel: (0273) 727328 Fax: (0273) 203754 Especially Fish / Amphibian / Reptile Books Also New Specialist Books, Publishers ’ Distributor We Accept Access, Visa, MC, EC, Am ex & JCB Please write stating interests Page 1 10 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 THE EIGHTH ANNUAL MIDWESTERN HERPETOLOGICAL SYMPOSIUM OCTOBER 16th - 18th, 1992 sponsored by the Wisconsin Herpetoiogical Society in celebration of their 20th Anniversary SCHEDULE OF EVENTS Fridev. Oct. 16th 4:00 PM - 8:00 PM Registration 5:30 PM - 7:00 PM Set-up for Swap Meet 7:00 PM - 11:00 7 SWAP MEET 7:00 PM - 11:00 ? ICE BREAKER (in conjunction with Swap) Saturday. Oct. 17th 8:00 AM - 8:30 Donuts, Coffee, Cool Pop 8:30 AM - Noon Talks 1 :00 PM - 5:30 PM Talks 7:30 PM - 9:00 PM Banquet - Milk Snake Slide Show with Author, Kenneth Williams 9:00 PM - 10:30 Auction Sundav. Oct. 18th 8:00 AM - 11:00 AM Talks 1 1 :00 AM - 5:00 PM Zoo Tour (own transport) Midway Hotel 1005 S. Moorland Rd. Brookfield, Wl 53005 Exit Moorland Rd from I-S4 m o t aJ is just expressway. for reMrvstioR* ati: (414) 786-9540 A block of rooms will be held for Symposium attendees till Sept. 24. 1992 special room rates: Single $59/night Double $65/night More than two to a room is an additional $6 per person G.a.- 4 people ■= $65 + $6 + $8 « $77) Indoor pool, sauna, and whirlpool available to all registered guests. When making reservations you must mention tha Eighth Midwest Herp Symposium sponsored by tha Wise. Harp Soc.to receive reduced room rates. Also make aura to get a confirmation number to guarantee your room. REGISTRATION FOR SYMPOSIUM THE LINE-UP QP SPEAKERS Dale Bertram - Topic: to be announced Gary Casper - Update on Wisconsin's Herp Atlas Project David Chiazar - "Strike Induced Chemosensory Searching and Chemical Clues Used by Rattlesnakes During Predatory Episodes" Michael Goode - "Reproductive Patterns in Freshwater Turtles’ Robert Hay - Ornate Box Turtles in their Natural Habitat and in Captivity Robert Henderson - "Distribution and Geographical Variations in the Wide Spread Neo-tropicel Tree Boa Corai/us enydris " Tom R. Johnson - Illustrating the Peterson Field Guide John Meltzer - "Captive Reproduction and Husbandry of the Western Hognose Snake" Michael Miller - "When to Take Your Herp to the Vet", end File Snakes Chris Palmer - Husbandry and Reproduction of Amphibians David Sorensen - Husbandry and Reproduction of the Rough-scale Sand Boa Eryx ccnicus Peter Strimple - Asian Monitors Thcmaa Tyning - "Uncommon Behavior of Common Amphibians and Reptiles" Kenneth Williams - "Honduran Cloud Forest Herpetology," and a special Slide Show presentation during the Banquet on Milk Snakes Pius, motet! Number _____ Full Registration (Talks, Banquet! $52 _____ Late Registration (after Sept. 30) $67 _____ Talks only $35 _____ Talks only (late registration) $45 _ Students under 1 6 (talks only) $20 _____ Banquet only $20 _ _ Banquet only (late registration) $22 TOTAL _ _ s«nd to: Wisconsin Herpetoiogical Soc. P.O. Box 366 Germantown, Wi 53022 There'll be a swap meet for symposium attendees: Trophy to the Best of Show. Tables will be available at the swap meet tor a donation to the Auction. Only captive bred animals - no imports. For reservations ceii (414) 353-4375. No venomous animals allowed in or near Hotel. The selling of animals in hallways, parking lots, and other public areas is not permitted. Please bring your photographs of "People and Herps," preferably 8 x 10's, for our display board. Attendees will vote on their favoritel Bulletin of the Maryland Herpetoiogical Society Page 1 1 1 Volume 28 Number 3 September 1992 ?lEWS & 9{0TL$ Sea Turtle Restoration Project EARTH ISLAND INSTITUTE 300 BROADWAY SUITE 28 SAN FRANCISCO CA 94133-3312 USA Telephone: 415-788-3666 FAX: 415-788-7324 Telex: 6502829302 MCI UW MCI Mail: 282-9302 EcoNet: earlhisland EQR IMMEDIATE RELEASE CONTACT: Kathy Nielsen: 415-788-3666 or 800-859-SAVE (7283) EARTH ISLAND SPONSORS HANDS-ON EXPEDITIONS TO SAVE THE GIANT SEA TURTLES Volunteers to assist conservation projects on Pacific coasts of Costa Rica and Nicaragua San Francisco, June 24, 1992 — Earth Island Institute's Sea Turtle Restoration Project today announced its volunteer expeditions to three of the most important sea turtle nesting beaches for the endangered olive ridley as part of its continued worldwide efforts to protect and restore threatened sea turtle populations. The olive ridley is known for its massive "arribadas, " the near simultaneous arrival of tens of thousands of nesting sea turtles. "Arribadas" have been described as the most spectacular event in the reptile world. "Watching an ancient sea turtle drag herself out of the dark surf and lay her eggs as her kind has done for eons is an experience of a lifetime," said the Sea Turtle Restoration Project's Expedition Director Kathy Nielsen. "Seeing an 'arribada ' is an incredible event experienced by very few people, something even most sea turtle biologists have not seen in person.” These expeditions will travel to three of only twelve known beaches in the world where this fantastic phenomenon occurs. Volunteers will team up with Latin and North American ecologists and marine scientists to assist in studies to unravel the mysterious life histories of these ancient animals and protect nesting sites. "We believe that a successful eco-tourism venture must meet the needs of the Earth and the local people, " Nielsen commented. "Participants will be helping conservationists with work that is of critical importance for this species' survival." Page 1 12 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 Volunteers choose to work at one of three beaches: Playa Nancite, Costa Rica; Ostional, Costa Rica; or Playa La Flor, Nicaragua. All three beaches are surrounded by beautiful tropical marine and forest habitats offering incredible wildlife viewing opportunities of such exotic animals as crocodiles, dolphins, coatis, howler monkeys, parrots, and iguanas. In addition to the olive ridley, three other species of sea turtles nest on these beaches including the leatherback. Leatherbacks can grow to be over nine feet long and weigh more than 2,000 pounds. The Expeditions range in cost from $500 to $1250, averaging $1000, not including airfare. Because participants will actually be working as volunteers for the non-profit environmental organization. Earth Island Institute, most or all of the costs, including airfare, are tax-deductible. The trip lengths vary from 7 to 14 days, with most trips being 10 days long. Volunteers need no experience, but must be in good physical condition. People interested in the Sea Turtle Restoration Project Volunteer Expeditions should call 800-859-SAVE or 415-788-3666, or write to Sea Turtle Restoration Project, Earth Island Institute, 300 Broadway, Suite 28, San Francisco, CA 94133 for more information. ##### Bulletin of the Maryland Herpetological Society Page 113 Volume 28 Number 3 September 1992 THE NEW YORK TURTLE AND TORTOISE SOCIETY 163 Amsterdam Avenue, Suite 365 New York, NY 10023 (212) 459-4803 FOR IMMEDIATE RELEASE FOR ADDITIONAL INFORMATION CALL: Allen Salzberg (718) 275-3307 (718) 275-7445 NEW YORK- TORTOISE INTERNATIONAL CONFERENCE ON TORTOISE & TURTLE CONSERVATION TO BE HELD IN JULY 1993 Purchase, New York Event to be Sponsored by the American Museum of Natural History's Turtle Recovery Program and the New York Turtle & Tortoise Society New York, NY, July 21, 1992 — The American Museum of Natural History's Turtle Recovery Program and the New York Turtle & Tortoise Society announced today that they will co-sponsor Conservation, Restoration, and Management of Tortoises and Turtles — an International Conference. The conference will be hosted by the State University of New York, SUNY Purchase and held Jjuly ll-_ 17. 199 3 .,) The SUNY Campus is located in Westchester County, just 3 0 minutes north of New York City. The conference was made possible by a grant from the Bureau of Land Management (U.S. Department of the Interior) . The conference will provide a forum for a wide array of people concerned with conservation to critically assess current efforts and develop strategies for the future. Using turtles as a model, the conference’ will develop techniques that can be applied to the conservation of a wide array of flora & fauna. This "diversity of opinions" is necessary for the success of the conference because, according to Dr. Michael Klemens, the Turtle Recovery Program Director, and the originator of the conference's theme, "conservation today is a dynamic, multi-disciplinary field encompassing a variety of areas including science, economics, public policy, sociology, even philosophy." Conference participants will include representatives from the academic, business, government, and private sectors including scientists, students, conservationists, wildlife managers, economists, land-use planners, government officials and policy makers . -over- Dedicalcd to the Appreciation of Turtles and Tortoises— to their Consen-ation and Preservation of their Habitat, to the Promotion of Proper Husbandry and Captive Propagation, and to the Education of the Public Page 114 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 9&WS dr 9\COT£S PAGE 2 Suzanne Dohxn, President of the New York Turtle & Tortoise Society, predicts that ,fwe'll have representation from six continents . " The Turtle Recovery Program is a joint initiative of the American Museum of Natural History and the IUCN-World Conservation Union Species Survival Commission. The program designs and implements turtle conservation programs around the world. The New York Turtle & Tortoise Society, one of the world's largest turtle groups, is dedicated to the conservation and preservation of turtles and tortoises and their habitat. The society is also committed to the promotion of the proper husbandry and captive propagation of turtles find tortoises. Conference proceedings will be published. For further information on the conference write to Craig Vitamenti c/o The New York Turtle & Tortoise Society, 163 Amsterdam Avenue, Suite 365, New York, NY 10023. Or call (212) 459-4803. # # # # # Bulletin of the Maryland Herpetological Society Page 115 Volume 28 Number 3 September 1992 9&WS &9&FES Ik _ A Ik. A Do You Have A Computer Modem ? Harp -Met Access: (215) 464-3562 _ _ - is an open access computer bulletin board serving the national herpetological community. It contains areas where one can ask questions, exchange information, and discuss reptiles. There are many text files that can be downloaded to your computer, as well as a database of all national and local herp organizations in the LJ.S. IBM USERS: send a full-featured $ 5.00 for a copy of communications program ] USE ANY PC, ANY MODEM. Any computer or terminal can access one of § six lines at 300/1200/2400 bps, 24 hours a day at (215) 464-3562. is an independent, community supported project carried on Satronics TUBS multi-line hos t Send a SASE to P.O. Box 52261, Philadelphia, PA. 19115 for more information on Page 1 16 Bulletin of the Maryland Herpetological Society Volume 28 Number 3 September 1992 OfE/WS & 6 o 4) a m 4m* fig 5) •St a «) X) 3 c« C 3 o ! <% eg c o tn "C as a 6 o . o eg s'3 1 1 c5 -w 3 W C g CO « CSS 4) 2 £ N s o o < ^ co ^ 2 : o 5 rS °. « Q co »m :ru - - d~S|J8<5 111 i| i'MIi8* ^ CO 10 . cs < css Q o U & 4> 5 a a ~< Bn Q S * ®<2 d m * u 11 ^>8 £d * i) «P fd W AS 0* g T3 w s § oS | Cd . £ |Is D 9 CO g Cd g a Cd N a o z us j QQ gS z g m I oo co "V nr c 4) 2 o o o 0) S d o 4> ”0 e s co Q CO n ^ IO m IS CO o S o g 5 o S o Jo q ej ffl O Q - §«g8f «£l£gj w ~ 8 3 xO 1 i 1 fH 52.0% 1 ci LO II CQ II ii sa oo IS 0) N oo N 00 Cl Cl N is m 00 \ LO N a N CM 05 CO © 00 \ 05 CM tjj W 8 t | & S> & I 1 w f d cd 6 cd d cd X5 d cd d cd 0) 10 \ 00 1 6 cd Page 144 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 Fig. 8. Map showing location of the Mescalero Sand Dunes, with locality records in text, and the local distribution of their associated subspecies of Sceloporus. Bulletin of the Maryland Herpetologicai Society Page 145 Volume 28 Number 4 December 1992 Page 146 Bulletin of the Maryland Herpetological Society Map showing distribution of the four species and subspecies in the Sceloporus undulatus superspecies. Intergrade areas are not indicated. Volume 28 Number 4 December 1992 Literature Cited Applegarth, John S. 1969. The variation, distribution, and taxonomy of the eastern fence lizard Scelopoms undulatus Bose in Latreille, in northeastern New Mexico. M. S. Diss., 124 pp. University of New Mexico, Albuquerque. Axtell, Ralph W. 1 988. Scelopoms graciosus. Interpretive atlas of Texas lizards, (5) : 1-4, map 3. Bell, Edwin L. 1954a. A taxonomic and evolutionary study of the western fence lizard, Scelopoms occidentalism and its relationships to the eastern fence lizard, Scelopoms undulatus. Ph. D. Diss. 163 pp., Univ. of Illinois, Urbana. 1954b. A preliminary report on the subspecies of the western fence lizard, Scelopoms occidentalism and its relationships to the eastern fence lizard, Scelopoms undulatus . Herpetologica 10(1): 31-36. Cole, Charles J. 1963. Variation, distribution, and taxonomic status of the lizard, Scelopoms undulatus vtrgatus Smith. Copeia, 1963(2): 413- 425. 1975. Karyotype and systematic status of the sand dune lizard {Scelopoms graciosus arenicolous) of the American Southwest. Herpetologica, 31: 288-293, figs. 1-2. 1983. Specific status of the North American Fence lizards, Scelopoms undulatus and Scelopoms occidentalism with comments on chromosome variation. Am. Mus. Nov., (2768): 1-13, figs. 1-5. Collins, Joseph T. 1990. Standard common and current scientific names for North American amphibians and reptiles.Third edition. Soc. Study Arnph. Rept. Herp. Circ., (19): i-iii, 1-41. 1991. Viewpoint; a new taxonomic arrangement for some North American amphibians and reptiles. Herp. Rev., 22(2): 42- 43. Bulletin of the Maryland Herpetologica! Society Page 147 Volume 28 Number 4 December 1992 Degenhardt, William G, and Kirkland L. Jones. 1972. A new sagebrush lizard. Scelopoms graciosus , from New Mexico and Texas. Herpetologica, 28; 212-217, figs. 1-2. Ferguson, George McNeil. 1982. Distribution, variation, and phenetic relationships of the lizard Scelopoms cautus Smith in northeastern Mexico. Ph. D. Diss. 195 pp., Univ. Texas, El Paso. Frost, Darrel R. and David M. Hillis. 1 990. Species in concept and practice: herpetological applications. Herpetologica, 46: 87-104. Germaro, A. L. 1972. Home range and movements of Holbrookia metadata metadata in eastern New Mexico. Herpetologica, 28: 165-168, figs. 1- 3. Jackson, James F. 1973. Distribution and population phenetics of the Florida scrub lizard, Scelopoms woocli. Copeia, 1973(4): 746-761. Leviton, Alan E., Robert H. Gibbs, Jr., Elizabeth Heal and C. E. Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology. Copeia 1985 (3): 802-832. Lowe, Charles H., Jr. and Kenneth S. Norris. 1956. A subspecies of the lizard Scelopoms undulatus from the White Sands of New Mexico. Herpetologica, 12: 125-127. Maslin, T. Paul. 1956. Scelopoms undulatus erythrocheilus ssp, nov. (Reptilia, Iguanidae), from Colorado. Herpetologica, 12: 291-294. Mayr, Ernst and Peter D. Ashlock. 1991. Principles of systematic zoology. Second edition, New York, McGraw-Hill, xx + 475 pp. Rand, Matthew Scholz. 1991, Behavioral function and hoimonal control of polymorphic sexual coloration in the lizard Scelopoms undulatus erythrocheilus. Ph. D. Diss. 147 pp. Univ. of Colorado, Boulder. Page 148 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 Sinervo, Barry. 1990. The evolution of maternal investment in lizards; an experimental and comparative analysis of egg size and its effects on offspring performance. Evolution, 44(2): 279-294. _____ and Raymond B, Huey. 1 990. Allometric engineering: an experimental test of the causes of interpopulational differences in performance. Science, 248: 1106-1109. Smith, Hobart M. 1938. Remarks on the status of the subspecies of Sceloporus undulatus , with descriptions of a new species and subspecies of the undulatus group. Occ. Pap. Mus. Zool. Univ. Michigan (387): 1-17, map. . Matthew S. Rand, J. David Drew, Bruce D. Smith, David Chiszar, and Christine M. Dwyer. 1991. Relictual intergrades between the Northern prairie lizard ( Sceloporus undulatus garmani) and the Red-lipped plateau lizard (S. u. erythrocheilus) in Colorado. MW Nat., 72: 1-11, figs. 1-8. _ _ and Rozelia B. Smith. 1976. Synopsis of the herpetofauna of Mexico. Vol. III. North Bennington, Vermont, John Johnson. 997 pp. Vinegar, Marian B, 1972. The function of breeding coloration in the lizard, Sceloporus virgatus. Copeia, 1972(4): 660-664. Dept of Environmental, Population, and Organismic Biology, University of Colorado , Boulder, Colorado 80309-0334 (HMS); Dept of Biology, Albright College, Reading, Pennsylvania 19612-5234 (ELB); 3293 West 14th Street Eugene , Oregon 97402-3193 (JSA); Department of Psychology, University of Colorado, Boulder, Colorado 80309-0345 (DC). Received: 6 May 1992 Accepted: 30 July 1992 Bulletin of the Maryland Herpetological Society Page 149 Volume 28 Number 4 December 1992 NATURAL HISTORY NOTES ON A SMALL POPULATIONS OF ANOLIS BREVIROSTRIS (SAURIA:POLYCHRIDAE) FROM ALTERED HABITAT IN THE DOMINICAN REPUPLIC Jane A. Moster, Robert Powell*, John S. Parmerlee, Jr., Donald D. Smith, and Amy Lathrop In an altered habitat we examined aspects of natural history in a small population of Anolis brevirostris, a trunk anole endemic to Hispaniola. Males were significantly larger than females. Sex ratios favored males, but were not significantly different from an expected 1 : 1 ratio. Habitat preferences were for larger broadleaf trees, among which lizards moved freely, and of which the trunks and larger limbs were used most frequently. Members of this lowland population appeared to be thermal conformers. Three distinct types of behavior were observed in males and females in a variety of situations. Joint¬ stepping was not restricted to prey approach, but used in intraspecific interactions as well. Males demonstrated aggression and territoriality. Interspecific interactions were limited to a few aggressive encounters with green anoles, A. chlorocyanus. Both sit-and-wait strategies and joint-stepping were used in feeding, most of which involved prey (largely ants and beetles) on tree surfaces. Larger individuals did not select larger food items and no significant differences in prey selection existed between males and females. The nematode Skrjabinoptera leiocephabrumis reported for only the second time from an anole. Anolis brevirostris is endemic to Hispaniola and distributed over the south -central part of the island and on many satellite islets (Schwartz and Henderson, 1991). These lizards are associated primarily with xeric habitats, but can be found in semimesic to mesic forests. Anolis brevirostris has been described as a trunk dweller (Williams, 1983) and prefers larger trees in forested areas, but is also found on fence posts, vines, shrubs, and on the ground under palm logs, stacks of palm fronds, and in coconut piles (Arnold, 1980). Ruibal (1965), Moermond (1981), and Jenssen and Gladson (1984) described behavior. Previous field work took place in essentially natural habitats, however, and since A. brevirostris is among several species of Hispaniolan anoles that have adapted well to altered areas, we address aspects of natural history in a small population of A. brevirostris in a heavily- trafficked park near the Hotel Guarocuya in Barahona, Dominican Republic. Page 150 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 Methods Anolis brevirostris weimorei from Barahona, Barahona Province, Dominican Republic were studied over a thirteen day period in June 1991. The ca. 750 m2 study site is a heavily used parkland situated near the southern edge of the city, approximately 35 m from the beach and 40 m north of the Hotel Guarocuya parking lot. The area contained sixteen broadleaf and palm trees planted in rows. To facilitate accurate recording of locations and perch heights, each tree and major branch was numbered, and lines were drawn around each tree and branch at one meter intervals up to four meters. Other lizards in the study area Included ground-dwelling Leiocephalus schreibersii, Ameiva chrysolaema, Celestas costatus , largely arboreal Anolis cy botes, Anolis chlorocyanus, and Anolis olssonu a grass anole. Individuals were noosed, sexed, measured, uniquely marked with paint to allow rapid recognition without resorting to recaptures, and toe- clipped for permanent identification. Cloacal temperatures and those of substrate and air at 2 cm and 1 m were taken with a cloacal quick-reading thermometer (Miller and Weber, Inc., Queens, New York). Environmental temperatures were taken in the shade and sheltered from wind. Population size was estimat ed using the Lincoki-Peterson index (Pacala and Roughgarden, 1985). The Schnabel method (Tanner, 1988) was used to evaluate if most or all of the animals were caught. Because these anoles are arboreal, ecological density (per tree) was computed in addition to crude density (by unit area). Lizards seen on branches extending outside of the study site boundaries were included if the tree trunk was within the plot. Seventeen specimens were collected outside the study site and preserved for stomach content analysis. Two additional specimens, collected in Barahona in May 1989, were also examined. Stomachs were excised and ingested food items were counted and identified to order. Volumes of contents were measured using methods of Milstead (1957), and importance values of food items were calculated according to Powell et al. ( 1990b). All means are presented plus or minus one standard deviation, and for all statistical tests, a < 0.05. Results and Discussion SIZE AND SEX RATIO.— Snout -vent lengths (SVL) of 47 adult males (41-52 mm, x=48.5±2.4 mm) were significantly longer than those of 33 adult females (38-46 mm, x=41.4±2.1 mm) (DF-78, t=-!3.7, P<0.001). The sex ratio (M:F) in the focal population was 33:27, that for the 19 lizards used in the investigation of food habits was 14:5. Neither the observed sex ratios in Bulletin of the Maryland Herpetologicai Society Page 151 Volume 28 Number 4 December 1992 the focal population (DF=3, x2-0.3Q, P-0.96), the anoles taken for stomach analysis (DF=3, x2=2„26, P=0.52), nor both combined (DF=3, x2=1.44, P=Q.70) differed significantly from an expected 1:1 ratio. The higher (but statistically insignificant) number of males may be explained by sampling error, as males of most iguanian species (sensu Frost and Etheridge, 1989) are likely to be more frequently encountered than females (Carpenter, 1 967) . This was most obvious in the sample taken for stomach content analysis, when the first 17 animals seen were captured, and no efforts were made to collect a representative sample or all of the lizards in a given area. POPULATION SIZE AND DENSITY.-— The focal population size was estimated as 57±1.77 (Lincoln-Peterson index). This does not account for immigration or emigration, both of which may have occurred. The Schnabel method indicated actual population size was approached. The mean number of lizards per tree was 3.6 and the crude density per unit area was 1 lizard / 13 m2 (-800/ ha). Although comparisons with populations in more natural locations are not available, these data indicate that A. brevirostris is capable of attaining high population densities in altered areas with considerable human activity. HABITAT UTILIZATION.-— Home ranges were not computed because of the difficulties associated with animals in three-dimensional habitats. Fitch et al. (1989) addressed this issue by calculating cylindrical areas used by trunk- dwelling Anolis cristatellus , but A. brevirostris often forages high into trees (despite their categorization as trunk- dwellers) where the number of small branches precluded accurate measurements of all substrates utilized. Furthermore, lizards readily moved from tree to tree, three instances of which were observed. In all, of 43 animals resighted at least once, 19 were observed on two different trees, five were seen on three different trees, and two were observed on four separate trees. Seventeen were seen only on the same tree. Eighteen individuals were not observed again after their initial capture. Lizards were found on all three of the larger broadleaf species in the area ( Aleurites molucana. Cassia siamea , Terminalia catappa} and moved between them freely; only 8 individuals (and almost always juveniles) were found on palms or smaller broadleaf trees. We recorded perch heights for 177 observations during which the lizard was seen before it moved (multiple observations of any individual were included only if from different days). Perch heights of adult males (N=125, 0. 1-4.5 m, x=2.1±l.l m) were significantly greater than those of adult females (N=46, 0. 1-3.2 m, x=1.4±0.9 m) (Mann-Whitney U, Z=-3.62, PcO.OOl). On only two occasions did we see individuals on the ground, but grit found among stomach contents of two animals suggested that these lizards, at least occasionally, forage on the Page 152 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 ground. Other trunk-dwelling anoles, A . cy botes and A. distichus , were observed to forage on the ground far more frequently in habitats (montane areas and confined sites, such as courtyards) in which ground- dwelling Ameiva and Leiocephalus were less common (or absent) than in the study area. Though quantitative data are lacking, we believe time spent on the ground by these anoles may be restricted by the presence of other, potentially competing species. THERMAL REGIME.— Cloacal temperatures (N=54, 28.2-32. 6°C, x-30.6 ± 3.4°C) were significantly correlated with substrate temperatures (N=45, 25.6-31 .8°C,x=28. 7 ± 3.2°C; N=39, r=0.43, PcO.Ol), air temperatures at 2 cm (N=50, 25.2-32.2°C, x=28.7± 1.7°G; N=44, r=0.61, PcO.OOl), and air temperatures at 1 m (N=50, 25.0-32.3°C, x"=28.6±1.5°C; N=44, r=0.37, P<0.02). Comparisons are given in Figure 1. Cole (1943) stated that substrate temperatures are more relevant to body temperatures than are air temperatures, because heat is primarily absorbed from the sun and the substrate. However, P. E. Hertz (pers. comm.) stated that substrate temperatures are not relevant to a small arboreal lizard, but the higher correlation in this study between cloacal and substrate temperatures than with air temperatures indicated that at least some heat exchange occurred with the substrate. Well-defined basking behavior was not observed. Most observations (94/116, 81%) were of individuals in shade (lizards often followed the shadows as they made their way around a tree in the course of a day), rarely (22/116, 19%) in diffuse sunlight, even during early morning or evening periods. Individuals were observed in direct sunlight only when disturbed by human activities. Activity periods extended from dawn until dusk. The combination of consistently warm weather, low elevation, and a largely shaded area seemed to provide suitable temperatures for optimal activity without the need for active thermoregulatory behavior. Hertz and Huey (1981) observed similar situations in their study of other species of Hispaniolan Anolis. BEHAVIOR. — Display behaviors of A. brevirostris were described by Jenssen and Gladson (1984), who defined type-A behavior as 5-7 bobs often followed by dewlapping. Bobs are presumably the push-ups of Ruibal (1965), who made a distinction between these actions. Because type-A behavior is common, and we only observed 6 instances of head bobbing (as described by Ruibal, 1965), we follow Jenssen and Gladson (1984) in not distinguishing between the two. Type-A displays are not always directed and may be used by patrolling males or employed after eating, defecating, or in copula. Type-B behavior, restricted to definite intraspecific interactions by Jenssen and Gladson (1984), was described as a series of bobs (= push-ups Bulletin of the Maryland Herpetological Society Page 153 Volume 28 Number 4 December 1992 of Ruibal, 1965) with simultaneous dewlapping. Type-C behavior is described here because observed behaviors did not all fall into previously defined categories. This behavior included tail-lashing and upraising (except during defecation) and is most often seen in combination with other behavior types (Jenssen and Gladson, 1984). Joint-stepping was similar to advances described by Moermond ( 198 1) in association with feeding behavior. Of 156 total field observations of behavior, 91 were characterized as type -A behavior, 31 involved dewlapping, 17 push-ups, and 43 push-ups followed by dewlapping by either one or more lizards in a given encounter. Although Jenssen and Gladson (1984) did not observe type -A behavior in females, in this study two female A. brevirostris were seen bobbing during copulation. Also, while Jenssen and Gladson (1984) included tail upraising in Anolis caudalis type -A behavior, but not in that of A. brevirostris , we observed tail upraising during intraspecific interactions five times in the course of this study. Seven different males displayed type-B behavior, on one occasion repeatedly. Two of these incidents also involved tail-lashing. Type-C behavior, alone or in combination with other behaviors, was observed on ten separate occasions by eight different individuals, only one a female (during pursuit by a male). Six of these incidents involved tail-lashing (always by males), and on two occasions (same individual) followed by tail- upraising. All observed type-C behaviors were directed. We observed joint- stepping separate from feeding behavior a total of 24 times, always involving cautious approaches or avoidance of other individuals. Many of the male-male interactions were categorized as aggressive, based on observations of type-A or B behavior, chasing, and/or biting. Though actual intent could not be determined, 21 incidents appeared to involve protection of a territory or a female within the area being protected, usually a branch or group of branches at nearly the same height. Four instances of aggressive male-male interactions were the result of intentional introductions of males into the vicinity of another male. In one case no aggressive action occurred. On the other three occasions, the introduced male was chased upward. None of the introduced males displayed any aggressive behavior, but all three of the resident males exhibited both type- A and B behavior during the encounters. Because chases are so rapid, it was difficult to observe biting action, although in one instance an introduced male was bitten three times while being chased. This individual took sanctuary on a terminal leaf. The resident male attempted an approach but failed. This was the only observation of an A. brevirostris on a leaf. Two of three introductions of females onto distant trees resulted in apparent acceptance by resident males, on the other occasion the introduced female Page 154 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 was chased and bitten twice by a male. No incidents of female-female aggression were observed. Six instances of displacement were observed. One was a male-male interaction in which an aggressor chased another from its perch, then took its place . The other five incidents were male-female interactions in which the male would move, followed by the female taking his spot. No apparent aggression was evident in these encounters. Williams (1983) described the primary escape behavior of trunk anoles as squirreling, upon disturbance lizards initially move to the opposite side of a branch or trunk, but will take refuge higher in the tree if provoked further. We observed escape behavior as a result of intra- and interspecific interactions, and in response to disturbance by humans. Most individuals reacted proportionately to the threat posed, squirreling, then crawling up the tree or out on limbs in increments upon being approached. In 4 1 of 46 recorded incidents involving responses to human observers, lizards moved upward and/or outward (four moved to different branches and one to a different tree), in 10 cases movement was accompanied by type -A behavior. On no occasions involving humans was a downward escape observed. In the five incidents during which lizards remained stationary (one turned to face upwards but did not ascend), three involved some form of type -A behavior. Of 22 male-male interactions recorded, nine involved chases of which eight concluded in upward or outward escapes. Four of these encounters involved type -A behavior. On one occasion a male Anolis chlorocyarius, a species usually observed at greater heights than A. brevirostris , chased an A. brevirostris upwards and bit it as it approached a female A. chlorocyarius. Near, but outside, the study area another male A. chlorocyarius was observed chasing a male A. brevirostris down the trunk, apparently the threat sufficient to overcome the lizard’s more commonly observed reluctance to escape downward. In addition to A. chlorocyarius , A. ay botes were found syntopically with A. brevirostris , but other than the incidents described above, no interspecific interactions were observed. Observations of reproductive behavior, in addition to displays and pursuits described above, included five incidents of copulation. In four cases lizards were facing down, in the other case orientation was not recorded. Twice the male was on the female’s right side, once on her left. Positions were not noted in the other two incidents. During copulation, males demonstrated combinations of type-A (N=5) and C (N=2) behavior. Type-A behavior was also observed in two females, although Jenssen and Gladson (1984) did not associate this behavior with females in copula. When finished, the pairs abruptly separated, whereupon the female ran up the tree in three instances and remained still in another sighting. Male post- Bulletin of the Maryland Herpetological Society Page 155 Volume 28 Number 4 December 1992 copulatoiy behavior included type-A behavior on two occasions, one of which also involved type-C behavior (push-ups with the tail upraised). FOOD HABITS.— -Of 24 recorded observations of prey capture, 16 involved a sit-and-wait strategy and in eight instances joint-stepping was noted. Moermond (1981) described advances, herein called joint-stepping, as a prey attack behavior of trunk anoles (although, as noted above, we saw it utilized more frequently in cautious approaches to other individuals). In 20 sightings, A. brevirostris captured and consumed prey on the tree, most likely ants (judging from both stomach analysis and the abundance of ants seen on the trees). Three times lizards were also seen attempting to capture flying insects (none successfully). On two occasions lizards were seen walking or jumping backwards in order to catch an insect. Analysis of stomach contents (Figure 2) indicated that ants and beetles were the most important food items. One stomach contained plant material and 2 contained grit, most likely from adventitious ingestion during prey capture, in the latter case probably on the ground. Foraging on the ground is not uncommon in trunk anoles (Rand, 1962; Williams, 1983), although we observed no such incidents. Surprisingly, as larger individuals might be expected to consume larger food items, mean size of food items decreased slightly, although not significantly, with increased SVL (N= 19; r=-0.09, P=0.37). Importance values of different food items did not differ significantly between sexes (DF=12, t=0.002, P=0.99). PARASITISM. — - Skrj abinoptera leiocephalorum (Nematoda; Physalopteroidea) previously had been found only in Leiocephalus spp. (Greve and Powell, 1989; Powell et al., 1990a, c). Two male Skrj abinoptera leiocephalorum were found in stomachs of A. brevirostris , admixed with ingesta. In a study synchronous with this one (Fobes et al., 1992), Skrj abinoptera sp. (but probably S. leiocephalorum) were also found in A. cybotes from a nearby site in Barahona. Prevalence was 0.16 (3 of 19 stomachs examined). Intensities were one in both host specimens. Acknowledgments Janet R. Lynxwiler, Julia W. Smith, Peter A. Zani, Paul T. Schell, Michael C. Schreiber, Timothy M. Fobes, Lisa R. White, Hoa T. Bui, Jaime D. Villa, Scott A. Maxey, and Sascha Oerter helped in the field. Thomas A. Zanoni, Jardin Botanico Nacional, identified plants of the area. John H. Greve, Iowa State University, identified the nematodes. Sixto and Yvonne Inchaustegui, Universidad Autonoma de Santo Domingo, and Jose A. Ottenwalder, Parque Zoologico Nacional, facilitated opportunities for field Page 156 Bulletin of the Maryland Herpetologica! Society Volume 28 Number 4 December 1992 Fig. 1 . Temperature data for Anolis breuirostris from Barahona, Dominican Republic: mean cloacal (open circles), substrate (squares), and air (solid circles) temperatures during morning (0500-1100 hrs) (N=13), midday (1100-1600 hrs) (N=14), and evening (1600-2100 hrs) (N=21) periods. Bulletin of the Maryland Herpetological Society Page 157 Volume 28 Number 4 December 1992 ARACHNIDA (ARANEAE) COLEOPTERA DIPTERA HOMOPTERA HYMENOPTERA (FORMICSDAE) HYMENOPTERA (OTHER) LEP1DOPTERA (LARVA) G DO MATA UNIDENTIFIED ARTHROPOD PARTS PLANT MATERIAL GRIT 0 10 20 ARACHNIDA (ARANEAE) COLEOPTERA DIPTERA HEM1PTERA HOMOPTERA HYMENOPTERA (FORMiCIDAE) HYMENOPTERA (OTHER) 4 1 LEPIDOPTERA - 3 LEPIDOPTERA (LARVA) - 1 ODONATA - 2 0 B 200 100 0.142 ARACHNIDA (ARANEAE) COLEOPTERA DIPTERA HEfeSPTERA HOMOPTERA HYMENOPTERA (FORMICIDAE) HYMENOPTERA (OTHER) LEPIDOPTERA LEPIDOPTERA ODONATA UNIDENTIFIED ARTHROPOD PARTS PLANT MATERIAL GRIT 0.092 Fig. 2. Stomach contents of Anolis brevirostris {N=19) from the Dominican Republic: (A) frequencies of occurrence of each item (numbers of stomachs in which each item was found): (B) total numbers of each item; (C) total volume of each item in mm3. Page 158 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 work. Permits were graciously provided by Emilio A. Bautista M.# Depart amento de Vida Silvestre. This investigation was supported in part by Grant No. BBS-9100410 from the National Science Foundation awarded to RP. Literature Cited Arnold, D. L. 1 980. Geographic variation in Anolis hrevirostris (Sarnia: Iguanidae) in Hispaniola. Breviora (481): 1-31. Carpenter, C. C. 1967. Aggression and social structure of iguanid lizards, p. 87- 105. In W. W. Milstead (ed.). Lizard Ecology: A Symposium. Univ. Missouri Press, Columbia. Cole, L. C. 1943. Experiments on tolerance of high temperatures in lizards with reference to adaptive coloration. Ecology 24(1) :94- 108. Fitch, H. S., R. W. Henderson, and H. Guarisco. 1989. Aspects of the ecology of an introduced anole: Anolis cristatellus in the Dominican Republic. Amphibia-Reptilia 10(3):307-320. Fobes, T. M., R. Powell, J. S. Parmerlee, Jr., A. Lathrop, and D. D. Smith. 1 992. Natural history of Anolis cy botes (Sauria: Polychridae) from altered habitat in Barahona, Dominican Republic. Carib. J. Sci. 28(3-4): in press. Frost, D. R, and R. Etheridge. 1989. A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata). Univ. Kansas Mus. Nat. Hist. Misc. Publ. (81): 1-65. Greve, J. H., and R Powell. 1989. Skrjabinoptera leiocephalorum n. sp. (Nematoda: Physalopteroidea) in Leiocephalus spp. (Reptilia: Iguanidae) from Hispaniola. J. Parasitol. 75(4):677-679. Bulletin of the Maryland Herpetological Society Page 159 Volume 28 Number 4 December 1992 Hertz, P. E., and R. B, Huey. 1981. Compensation for altitudinal changes in the thermal environment by some Anolis lizards on Hispaniola. Ecology 62(31:515-521. Jenssen, T. A., and N. L. Gladson. 1984. A comparative display analysis of the Anolis brevirostris complex in Haiti. J. Herpetol. 18(31:217-230. Milstead, W. W. 1957. Some aspects of competition in natural populations of whiptail lizards (Genus Cnemidophorus) . Texas J. Sci. 9(4):410-447. Moermond, T. C. 1981. Prey- attack behavior of Anolis lizards. Z. Tierpsychol. 56:128-136. Pacala, S. W., and J. Roughgarden. 1985. Population experiments with the Anolis lizards of St. Maarten and St. Eustatius. Ecology 66(1): 129-141. Powell, R., P. J. Hall, and J. H. Greve. 1990a. Occurrence of Skrjabinoptera leiocephalorum (Spirurida: Physalopteridae) in Leiocephalus spp. (Sauria: Iguanidae) from Hispaniola. J. Helminthol. Soc. Washington 57(1):75- 77. _ . J. S. Parmerlee, Jr., M. A. Rice, and D. D. Smith. 1990b. Ecological observations of Hemidactylus brookii haiiianus Meerwarth (Sauria: Gekkonidae) from Hispaniola. Carib. J. Sci. 26(1-21:67-70. _ ., D. J. Pflanz, J. H. Greve, and D. D. Smith. 1990c. Life history notes (parasitism): Leiocephalus semilineatus. Herpetol. Rev. 21(3):60-61. Rand, A. S. 1962. Notes on Hispaniolan herpetology 5. The natural histoiy of three sympatric species of Anolis. Breviora (154): 1-15. Ruibal, R. 1965. Evolution and behavior in West Indian anoles, p. 1 16-140. In W. W. Milstead (Ed.), Lizard Ecology: A Symposium. Univ. Missouri Press, Columbia. Page 160 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 Schwartz, A. , and R. W. Henderson. 1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History. University of Florida Press, Gainesville. Tanner, J. T. 1 988. Guide to the Study of Animal Populations. Univ. Tennessee Press, Knoxville. Williams, E. E. 1983. Ecomorphs, faunas, island size, and diverse end points in island radiations of Anolis, p. 326-378. In R. B. Huey, E. R. Pianka, and T. W. Schoener (Eds.). Lizard Ecology: Studies of a Model Organism. Harvard University Press, Cambridge, Massachusetts. Department of Biology, Saint Mary College, Leavenworth Kansas 66048 (JAM); Department of Natural Sciences, Avila College, Kansas City, Missouri 64145 ( RP * JSP); Division of Allergy & Rheumatology, University of Kansas Medical Center, Kansas City, Kansas 66103 (DDS); and Museum of Natural History , University of Kansas, Lawrence, Kansas 66045 (AL). * Corresponding author. Received: 20 May 1992 Bulletin of the Maryland Herpetological Society Page 161 Volume 28 Number 4 December 1992 BOELLA TENELLA IS EPICRATES 1NORNATUS (REPTfUA: SERPENIES) Van Wallach and Hobart M. Smith The holotype of Boella tenella Smith and Chiszar (1992), supposed to be from the state of Oaxaca, Mexico, actually represents Epicrates inomatus (Reinhardt) and is accordingly from Puerto Rico. Erroneous or unknown locality data on type material for newly described onomotaxa (nominal taxa) have plagued herpetological research from the beginning of taxonomic time (Linnaeus, 1758) to the present. Many such taxa, to be sure, are rather promptly placed with their proper biotaxa, resulting in no long-enduring problems. Examples are: Anilius scytale (Linnaeus, 1758) of northern South America (type locality “Indiis”) ; Conopsis nosusGunther(1858) of central Mexico (type locality “California); Drymobius margaritiferus (Schlegel, 1837) of Texas, Mexico and Central America (type locality “New Orleans”); Epicrates fordii (Gunther, 1861) of Hispaniola (type locality “western Africa”); Helicops angulatus (Linnaeus, 1758) of northern South America (type locality “Indiis”); Helicops trivittatus (Gray, 1849) of Brasil (type locality “India”); Hypnale nepa (Laurenti, 1768) of Sri Lanka (type locality “America”): Lamprophis aurora (Linnaeus, 1758) of South Africa (type locality “America”); L. Juliginosus (Boie, 1827) of Africa (type locality “Java”); Leptodeira maculata (Hallowell, 1861) of western Mexico (type locality “Tahiti”); L. punctata (Peters, 1866), also of western Mexico (type locality “South Africa”); Liophis triscalis (Linnaeus, 1758) of Cpragao (type locality “Indiis”); and Sibon nebulatus (Linnaeus, 1758) of Mexico, Central America and northern South America (type locality “Africa”). Other names proposed with no or incorrect locality data have posed lingering problems, especially when nomenclatural stability is threatened, as happens if the biotaxa involved had already acquired names, sometimes much used, of later date. Examples within our personal experience, although not significantly disruptive (i.e., not enough to justify an appeal to the International Commission on Zoological Nomenclature (ICZN) for name- conservation, include Glauconia boettgeri Werner (1899), Eumeces capita Bocourt (1879), and Scolecophis fumiceps Cope (1861). The first was accompanied by no locality data whatever (although subsequent speculation focused on Africa), Bocourt’s name by the published data “la cote oriental des Etats-Unis”, although its museum registers record “Mexique” and Page 162 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 “Ameiique septentrionale”, and Cope’s name by “probably Cuba.” Werner’s name ultimately proved to antedate Leptotyphlops humUis slevini Klauber (1931) of the Cape region of Baja California; Bocourt’s name (Smith, Smith and Guibe, 1941) to antedate Eumeces xarithi Gunther ( 1 889) of China; and Cope’s name (Smith, 1941) to antedate TantUla kimia Blanchard (1938) of southern Texas and northern Tamaulipas. Since no appeal for name- conservation was considered to be warranted in these cases, the three taxa are now known as L. h» boettgert E. capita , and T. nigrtceps fumiceps, respectively. Appeal for conservation was made, successfully, in another case, involving Henicognathus sumichrasti Bocourt (1886), supposedly from “Cacopriets” (=Cacoprieto), Oaxaca, Mexico. Guibe and Roux-Esteve (1962) showed that the holotype is an example of the long-recognized Ablates chinensis Gunther (1889) of China (now Sibynoptus c. chtnensis). Bocourt’s earlier name was suppressed, and Gunther’s name conserved, by the ICZN in 1965. Generally, however, onomotaxa proposed for type material with incorrect or no locality data prove ultimately to have been applied to biotaxa having been given names previously. Thus such erroneously created names sink into oblivion as junior synonyms, never to be revived unless their senior synonyms are discovered to be invalid for one reason or another (although such jr. synonyms ordinarily are forever invalid, the names are forever occupied and cannot be used later even in a different context, in the same genus). Numerous such cases are known. Werner (1924, 1925) proposed thirteen new snake genera and species based on type material with no or incorrect locality data, all but two of which are now known to be jr. synonyms of earlier names (Wallach, 1988). Other examples include Cope’s (1861) Prymnomiodon chalceus , supposedly from “Siam”, which proved to be a jr. synonym of Eutaenia sackenii Kennicott (1859) (now ThamnophLs sauritus sackenir, Rossman, 1961) of Florida ; and Malnate’s (1968) NatrixdunnU type locality “Kuala Lumpur, Malaysia”, is a jr. synonym of ThamnophLs cyrtopsis (Kennicott, 1860) fide Rossman (in litt.). Muller’s (1923) Cochliophagus tomieri from “South America”, is ajr. synonym of Sibonfasciatus (Gunther, 1858) of Mexico, fide Amaral (1929). Similarly, Fischer’s (1885) Leptognathus alhocinctus, from “San Francisco, California”, is ajr. synonym (Kofron, 1985) of Sibon phUippi (Jan, 1863), of western Mexico. To the list in the preceding paragraph (which could be considerably extended, as could the others) we here add Boella tenella Smith and Chiszar (1992), supposedly from Oaxaca, Mexico, but which, as determined initially by VW, actually represents Epicrates inomatus (Reinhardt, 1843) of Puerto Rico. For the egregious error HMS assumes full responsibility. Curiously, Bulletin of the Maryland Herpetoiogical Society Page 163 Volume 28 Number 4 December 1992 fide Schwartz and Thomas (1975), Boella is the second jr. synonym acquired by E. inomatus based on material with erroneous locality data; the first is Piesigaster boettgeri Seoane (1881), supposedly from “Mindanao, Philippine Islands.” In examining the holotype of Boella tenella (UCM 56508), VW discovered that numerous major errors of anatomical interpretation appeared in the original description, contributing in part to the incorrect taxonomic conclusions. Contrary to the statements there, the hyoid cornua are divergent (and possibly joined anteriorly), hypapophyses are absent on the posterior vertebrae, and the specimen is a juvenile male. Additionally, there is no tracheal lung, a large left lung is present, and the tracheal entry into both left and right lungs is subterminal. All of these characters demonstrate that the specimen is not a member of the Tropidophiidae but of the Boidae (McDowell, 1987). Among the Boidae, the specimen can undoubtedly be allocated to Epicrates. A visceral structure peculiar to Epicrates is the sudden constriction of the left lung, leaving the posterior half greatly reduced in diameter to a thin tail. The left lung of Boella exhibits a thin tail along the caudal 52% of its length. The fact that the specimen is a male is revealed by the presence of an immature testis and paired hemipenes. A peculiarity in the viscera of this specimen is indicated by the apparent presence of only one undeveloped testis (right, 1.1% SVL) and one adrenal gland (right, 2.1 % SVL), both of which are located adjacent to the right kidney (right testis midpoint 73.5%, right adrenal midpoint 74.1%, and right kidney midpoint 74.1% SVL), a most unusual situation for any snake. The gonads and adrenals are located craniad of the kidneys in most snakes although in some species the caudal tip of the left gonad may overlap the cranial tip of the right kidney (Wallach, 1991). In support of the fact that the specimen is a juvenile, the testis is underdeveloped (1% SVL; adult Epicrates have testes 3-6% SVL) and an umbilical scar (four ventrals in length) is present. Sheplan and Schwartz in 1974 were not aware of any young of this species in American collections. Within Epicrates the type of Boella keys out to be an Epicrates inomatus of Puerto Rico (Schwartz and Henderson, 1985). Additionally, all of its scale counts (except subcaudals, which lower the range for the species by one) fall within the range of variation of E. inomatus as reported by Sheplan and Schwartz (1974). Following are the counts of Boella with the ranges reported by Sheplan and Schwartz for E. inomatus (male counts only for ventrals and subcaudals) given in parenthesis: ventrals, 264 (263-273); subcaudals, 65 (66-74); ventrals plus subcaudals, 329 (329-338); anterior Page 164 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 scale rows, 33 (31-34); midbody scale rows, 39 (38-42); posterior scale rows, 21 (21-25); supralabials, 11-12(11-1 2); supralabials entering orbit, 6-7 (5- 7); infralabials, 12-13 (11-15); loreals, 2 (1-2); circumorbitals, 8-9 (7-10); pre-intersupraoculars, 2 (2-3); intersupraoculars , 1 (1); and post- intersupraoculars, 2 (2-3). Stejneger’s (1904) figure of the head scales of a young specimen (USNM 12446) is very similar to the Boella type. The greatest peculiarity of the latter is the fusion of intemasals and nasals; the combined scales were interpreted as nasals in contact medially by Smith and Chiszar (1992), a condition apparently not recorded previously in Epicrates. That interpretation is presumably untenable, the nasals being fused anomalously with the intemasals. A comparison of the internal anatomy of Boella with that of a subadult female (770 mm SVL) Epicrates inomatus (UMMZ 74415) reveals a remarkable agreement in morphology, placement, and size of the viscera, especially when considering normal ontogenetic and sexual variation (WaEach, 1991). Following are the visceral data, expressed as % snout-vent length (SVL) unless otherwise indicated, for Boella and (parenthetically) Epicrates inomatus : posterior tips of hyoid, 6.5 (5.2); trachea plus intrapulmonaiy bronchus length, 32.2 (32.0); trachea plus intrap ulmonary bronchus midpoint, 16.1 (16.0); approximate number of tracheal rings, 419 (434); width of tracheal membrane /width of tracheal rings, 0.3 (0.2); heart length, 2.3 (2.6); heart midpoint, 27.3 (27.9); junction of systemic arches, 29.9 (29.2); orifice of right lung (midpoint), 29.5 (29.9); cartilages in right intrapulmonaiy bronchus, 5 (11); right lung length, 28.9 (30.3); right lung midpoint, 41.6 (44.0); right lung posterior tip, 56. 1 (59. 1); left lung length, 19.3(19.5); left lung midpoint, 37.2 (38.8); left lung posteriortip, 46.9 (48.6); left lung /right lung, 66.7 (64.4); heart-liver gap, 8.4 (6.8); liver length, 17.6 (18.7); liver midpoint, 45.6 (45.3); liver-gall bladder gap, 7.3 (6.8); liver-gall bladder interval, 26.8 (27.1); gall bladder midpoint, 62.7 (62.3); right kidney length, 3.4 (3.6); right kidney midpoint, 74. 1 (77.5); number of right kidney lobes 10(10); number of right renal arteries 2 (1); left kidney length, 3.6 (3.4); left kidney midpoint, 80.7 (81.8); number of left kidney lobes 8 (9); number of left renal arteries 1 (1); right plus left kidney length, 6.9 (7.0); kidney-vent interval, 27.6 (24.3); kidney-vent gap, 17.6 (16.5); and umbilical scar-anal plate interval in % ventrals), 30.7 (31.2). The differences evident in the preceding comparisons are not significant, and indeed are smaller than might be expected. There can be no doubt as to the correct identity of Boella tenella. We therefore remove Boella from the family Tropidophiidae and place Boella tenella in the synonymy of Epicrates inomatus (Reinhardt, 1843), family Boidae. Bulletin of the Maryland Herpetological Society Page 165 Volume 28 Number 4 December 1992 Acknowledgments We thank Drs. Shi-Kuei Wu (tJCM) and Arnold G. Kluge (UMMZ) for permission to dissect specimens in their care. Literature Cited Amaral, A. do. 1929. Estudos sobre ophidios neotropicos. XVIII. Lista remissiva dos ophidios de regiao neotropica. Mems. Inst. Butantan 4: i-viii, 129-271. Bocourt, M. F. 1879. Etudes surles reptiles. Miss. Sci. Mexique, Rech. Zool. , Livr. 1886. 6: 361-440, pis. 21-22, 22A-22D. Etudes surles reptiles. Miss. Sci. Mexique, Rech. Zool., Livr. 10: 593-664, pis. 36-41. Boie, F. 1827. Bemerkungen uber Merrem’s Versuch eines Systems der Amphibien. Ite. Lieferung. Ophidier. Isis von Oken 20(6): 508-566. Cope, E. D. 1861. Catalogue of the Colubridae in the museum of the Academy of Natural Sciences of Philadelphia. Part 3. Proc. Acad. Nat. Sci. Philadelphia 12: 553-566. Fischer, J. G. 1885. Ichthyologische und herpetologische Bemerkungen V. Herpetologische Bemerkungen, Jb. Hamb. Wiss. Anst. 1884: 82-119, pis. 3-4. Gray, J. E. 1849. Catalogue of the specimens of snakes in the collection of the British Museum. British Mus. (Nat. Hist.), London, xv, 125 pp. Guibe, J. and R. Roux-Esteve. 1963. The generic status of the snake Henicognathus sumichrasti Bocourt, 1886. Herpetologica 18(4): 268-269. Page 166 Bulletin of the Maryland Herpetologicai Society Volume 28 Number 4 December 1992 Gunther, A. C. L. G. 1858. Catalogue of colubiine snakes in the collection of the British Museum. London, Taylor & Francis, xvl, 281 pp. 1861. On a new species of the family Boidae. Proc. Zool. Soc. London, 1861: 142, pi. 23. 1889. Third contribution to our knowledge of reptiles and fishes from the Upper Yangtsze-Kiang. Ann. Mag. Nat. Hist. (6)4: 218-229. Hallowell, E. 1861. Report upon the Reptilia of the North Pacific Exploring Expedition, under the command of Capt. John Rogers, U.S.N. Proc. Acad. Nat. Sci. Philadelphia 12: 480-509 Jan, G. 1863. Elenco sistematico degli ofidi descritti e desegnati per l’iconografia generale. Milan, Lombardi, vii, 143 pp. Kennicott, R. 1859. Notes on Coluber calligaster of Say, and a description of new species of serpents in the collection of the North Western University of Evanston, Ill. Proc. Acad. Nat. Sci. Philadelphia 11:98-100. 1860. Descriptions of new species of North American serpents in the museum of the Smithsonian Institution, Washington. Proc. Acad. Nat. Sci. Philadelphia 12: 328-338. Klauber, L. M. 1931. Notes on the worm snakes of the southwest, with descriptions of two new subspecies. Trans. San Diego Soc. Nat. Hist. 6(23): 333-352. Kofron, C. P. 1985. Systematics of the neotropical gastropod-eating snake genera, Tropidodipsas and Sibon. J. Herp. 19(1): 84-92, figs. 1-3. Laurenti, J. N. 1768. An striae i viennensis specimen medicum. . .Joan. Thom. Nob. deTrattnem, Vienna. 241 pp., 5 pis. Bulletin of the Maryland Herpetological Society Page 167 Volume 28 Number 4 December 1992 Linnaeus, C. 1758. Systema naturae... Tenth ed. Vol. 1. Stockholm, Salvius. iv, 826 pp. Malnate, E. V. 1 968. Matrix dunnu a new species of water snake from Malayasia. Notulae Naturae (410): 1-5. McDowell, S. B. 1987. Systematics. In: R. A. Seigel et al.. Snakes...: 3-50, fig. 1. Muller, L. 1923. Uber neue Oder seltene mittel- und sudamerikanischen Amphibien und Reptilien. Mitt. Zool. Mus. Berlin 11(1): 77- 94. Peters, W. C. H. 1 866. Uber neue Amphibien des zoologischen Museum zu Berlin. Mbr. dt. Akad. Wiss. Berlin 1866: 86-94. Reinhardt, J. T. 1843. Beskrivelse ad nogle nye Slangearter. Danske Vid. Selsk. Afhandl. 10: 233-279. Rossman, D. A. 1961. Nomenclatorial status of the neotropical subspecies of the Schlegel, H. 1837. colubrid snake, Thamnophis sauritus. Notul. Nat. (340): 1- 2. Essai sur la physionomie des serpens. La Hague, Kips & Van Stuckum. 28, 251, 314, 606 pp., 27 pis., 3 maps. Schwartz, A. and R. W. Henderson. 1985. A guide to the identification of the amphibians and reptiles Seoane, V. L. 1881. of the West Indies exclusive of Hispaniola. Milwaukee Publ. Mus., Milwaukee. 165 pp. Neue Boiden-gattung und -Art von den Philippinen. Abh. Senckenb. Naturf. Ges. 12: 217-224. Page 168 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 Sheplan, B. R, and A. Schwartz. 1 974. Hispaniolan boas of the genus Epicrates (Serpentes, Boidae) Smith, H. M. 1941. and their Antillean relationships. Ann. Carnegie Mus. 45: 57-143, figs. 1-6. Synonymy of Tantilla nigriceps fumiceps . Copeia 1941(3): 112. __ _ » and D. Chiszar. 1992. A Mexican genus of tropidopheine snakes. Bull. Maryland Herp. Soc. 28: 19-28, figs. 1-5. __ , R B. Smith, and J. Guibe. 1975. The identity of Bocourt’s lizard, Eumeces capita. Gr. Basin Nat. 35(1): 109-112, figs. 1-4. _ , and K. Larsen. 1974. The name of the Baja California Cape wormsnake. Gr. Basin Nat. 34(2): 94-96. Stejneger, L. 1904. The herpetology of Porto Rico. Rep. U. S. Natl. Mus. 129: 549-724, figs. 1-197. Wallach, V. 1988. Status and redescription of the genus Padangia Werner, with comparative visceral data on Collorhabdium Smedley and other genera (Serpentes: Colubridae). Amph.-Rept. 9: 61-76, figs. 1-3. 1991. Comparative visceral topography of African colubrid snakes of the subfamilies Aparallactinae and Atractaspidinae. Master’s Thesis, Louisiana State Univ., Baton Rouge. 490 pp., figs. 1-144. Werner, F. 1899. Beschreibung einiger neuer Schlangen und Batrachier. Zool.Anz. 22:114-117. 1924. Neue oder wenig bekannte Schlangen aus dem Naturhistorischen Staatsmuseum in Wien. Sitzber. Akad. Wiss. Wien 133: 29-56, figs 1-9. Bulletin of the Maryland Herpetological Society Page 169 Volume 28 Number 4 December 1992 1925. Neue oder wenig bekannte Schlangen aus dem Wiener naturhistorischen Staatsmuseum (2. Teil). Ibid. 134: 45-66, figs. 1-4. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 (VW); and Department of Environmental, Population and Organismic Biology, University of Colorado, Boulder, Colorado 80309 0334 (HMS). Received: 5 June 1992 Accepted: 7 July 1992 Page 170 Bulletin of the Maryland Herpetological Society Volume 28 Number 4 December 1992 STRIPED PRAIRIE KIN0SNAKES, LAMPROPELTIS CALLIGASTER. FROM ILLINOIS The occurrence of aberrant color patterns in snakes is relatively common, descriptions of such are found throughout the literature (reviewed by Bechtel 1978). More specifically, striping has been found in a number of snake taxa. For example, Gloyd (1958) described abnormal striping in Crotalus scutulatus and Crotalus horridus. This report represents the second published account of striping in Lampropeltis calligaster. Ashton ( 1973) described a single striped individual that hatched from a clutch of 13 eggs, all other siblings were normally patterned. In April of 1991 in Effingham County a normally patterned adult female L. calligaster (SVU=790 mm) was captured. On June 15, 1991 the female laid 10 normal sized eggs, two eggs were infertile and were later discarded. The eggs were incubated on damp vermiculite at 24-28° C. On August 13, 1991 six of the eight remaining eggs hatched. Five of the six hatchlings showed various degrees of striping (Table 1). One of the striped hatchlings (hatchling 6 in table 1) had a severe “kink” approximately two head lengths behind the neck and died 36 days after hatching. The remaining two eggs were opened August 15 and each contained a fully developed snake. One had severe bone deformations and was striped and the other appeared normal though small and was normally patterned. Table 1 shows snout-vent length, stripe length, the percent snout- vent length of striping, and the width of striping in terms of the number of dorsal scales covered. The range of the percent striping was 7.4% - 70.0% of the snout-vent length with a mean percent of striping of 42. 1% and standard deviation 25.9%. While working on another project, I discovered a specimen of L. calligaster from Osage County Oklahoma with a continuous mid-dorsal stripe in the collection of the Museum of Natural History at the University of Kansas (KU 154482). The stripe covered 1.5 to 2.5 dorsal scales. The snake had a snout-vent length of 497 mm and a stripe length of 510 mm. Bulletin of the Maryland Herpetological Society Page 171 Volume 28 Number 4 ~ £ £ JS 5 2 CO 4) . *£ a 2 5 2 . ttf) S £ s - o ^ *d j2 0 g «-» Q 03 y ^ 73 *° g W v 5 -3 id 1 25 ^ Cfl ^ v- cn o 73^ gjjg P> W l°« ^|s ^ a,) K •*=-> (0 £ n9 o 5 'afi C a 4) a TE 3) toO C 2 o ■*«* cd X $ in ^ d ^ eo CO in in o in bULLETIN of the TJRacylanb ^fr f)erpeto!ogical Ooriety DEPARTMENT OF HERPETOLOGY THE NATURAL HISTORY SOCIETY OF MARYLAND, INC. MdHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League 31 MARCH 1993 VOLUME 29 NUMBER 1 BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY Volume 29 Number 1 March 1993 CONTENTS New Variational Extremes for Tantilla calamarina and a Locality Record Correction for Conophis vittatus victims (Reptilia: Serpentes) Hobart M. Smith, Oscar Flores Villela and David Chiszar ..... 1 Phylogenetic Relationships and Taxonomy of Colostetus mandelomm (Anura: Dendrobatidae), with Notes on Coloration, Natural History, and Description of the Tadpole Enrique La Marca. . . . . . . . . . ....4 Variation in Two species (G. bicolor, G. duellmcmi) or Mexican Earth Snakes ( Geophis ) Hobart M. Smith and Oscar Flores Villela . . . . 20 Observations on the Sensitivity to High Temperatures In Two Lizards Species ( Ameiva undulata and Sceloporus horridus) from Zacatepec, Morelos, Mexico Julio A. Lemos -Espinal, Royce E. Ballinger and Juan M. Javelly-Gurria . . . . . . . 24 The Range of the So-called Relictual Intergrades between the Lizards Sceloporus undulatus Garmani and S. il srythrocheilus Hobart M. Smith, David Chiszar, Emmett Evanoff and Jeffry B. Mitton . . . .....30 The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc . 2643 North Charles Street Baltimore, Maryland 21218 BULLETIN OF THE Volume 29 Number 1 March 1993 The Maryland Herpefologicai Society Department of Herpetology, Natural History Society of Mar/Sand, Inc. Executive Editor.., . ...Herbert S. Harris, Jr. Steering Committee Frank Groves Jerry D. Hardy, Jr. Herbert S. Harris, Jr. Library of Congress Catalog Card Number: 76-93458 Membership Rates Membership in the Maryland Herpetological Society is $16.00 per year. Foreign $20. 00 /year. Make all checks payable to the Natural History Society of Maryland, Inc. Meetings The third Wednesday of each month, 8:15 p.m. at the Natural Histoiy Society of Maryland (except May-August, third Saturday of each month, 8:00 a.m.j. The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 p.m. Volume 29 Number 1 March 1993 NEW VARIATIONAL EXTREMES FOR TANTILLA CALAMARINA AMD A LOCALITY RECORD CORRECTION FOR CONQPHIS V17TATUS VIDUUS (REPTILIAN SERPENTES) Hobart M. Smith, Oscar Flores Villela and David Chiszar A juvenile female Tanlilla calamarina Cope (MZFC-UNAM (Museo de Zoologia, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico) 809) from near El Platanito, on the Tepic-Jalcocotan highway in Nayarit, Mexico, municipality of Tepic, exhibits variational extremes not previously recorded for the species in the exhaustive reviews by Wilson and Meyer (1981) and Wilson (1988). The specimen is only 77 mm in total length (tail 8 mm) , shorter than any of 22 females of the species recorded by Wilson and Meyer (1981), with a range of 104-202 mm. Its condition is very good, the body nearly straight but the neck sharply bent to the right at a 180° angle, leaving the head parallel with the neck. Most character states conform with the norms for the species (Wilson and Meyer, 1981): six supraiabials, the fifth separated from parietal; postoculars and preoculars 1-1, the latter in contact with postserninasal; first infralabials separated; 129 ventrals. There are, however, only 22 subcaudals (vs 25-33 previously recorded). The pattern on the head is the same as described and illustrated by Wilson and Meyer (1981), but the pattern on body is more elaborate than previously described. The usual vertebral dark line is present, although narrow and confined to the median third or less of the median scale row, and the typical broad dark stripe occupies the adjacent parts of the 3rd and 4th scale rows, more extensive on the former. However, all other scale rows, except the 7th on each side, have a continuous, narrow, median dark line, lighter than the three primary lines, but still very distinct. All lines extend from neck to tail, except for the one on the 6th scale row, which from about midbody to tail is represented by a series of dim dark spots. Only the primary dark lines continue onto the tail. The secondary lines constitute merely an enhancement of the separate dark streaks on most scales, as shown in Fig. 2 of Wilson and Meyer (1981), although the result is strikingly distinctive. Because of the many basic similarities to other examples of the species, and the numerous records for the same area whence the present specimen came, there is no reason to suspect that the latter represents anything but a variant of T. calarnarincL Bulletin of the Maryland Herpetoiogicai Society Page 1 Volume 29 Number 1 March 1993 This specimen was previously reported (Flores et aL, 1991: 182) erroneously as Geagras redimitus Cope, which is actually limited, so far as known at present, to Oaxaca (state) (Wilson, 1987). This is not the only occasion on which these two closely similar species have been confused, as noted by Wilson and Meyer ( 198 1 : 2 1), who called attention to a record of G. redimitus from Michoacan that in fact pertains to T. calamarina. We take this opportunity to correct an erroneous record (Flores et aL , 1991: 1 77) of Conophis vittatus viduus Cope for Merida, Yucatan, Mexico, to Coniophanes imperialis clavatus (Peters). The specimen in question (MZFC- UNAM 3170) is typical of its taxon, having 19-17 scale rows; 125 ventrals; 77 subcaudals; alight, dark-edged line through upper part of eye; a dark line through upper part of supralabials, which are immaculate white below that line; a continuous, narrow, vertebral dark line; sides of body dark below a dark line along middle of 4th scale row; lateral edges of ventrals with a small, irregular dark spot. These character- states agree fully with the ranges of variation summarized by Bailey (1939: 35-41). Although the latter author had seen no specimens from Yucatan, many were available for Lee’s (1980: 65) review. The present specimen thus constitutes no range extension or morphological variation not previously recorded. Literature Cited Bailey, J. R. 1 939 . A systematic revision of the snakes of the genus Coniophanes. Pap. Michigan Acad. Sci. Arts Letters, 25 (2): 1-48, figs. 1- 5, pis. 1-3. Flores Villela, O. A., E. Hernandez Garcia and Adrian Nieto Montes de Oca. 1991. Catalogo de anfibios y reptiles del Museo de Zoologia, Facultad de Ciencias Universidad Nacional Autonoma de Mexico. Univ. Nac. Auton. Mexico, Fac. Ciencias. xiii, 222 PP- Lee, J. C. 1 980. An ecogeographic analysis of the herpetofauna of the Yucatan Peninsula. Univ. Kansas Mus. Nat. Hist. Misc. PubL, (67): i- iii, 1-75. Wilson, L. D. 1987. Page 2 Geagras , G. redimitus . Cat. Am. Amph. Kept., (430): 1-2, map. Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 1988. Tantilla calamarina Cat. Am. Amph. Kept., (433): 1-2, map. __ and J. R Meyer 1981. Systematics of the calamarina group of the colubrid snake genus Tantilla I Milwaukee Public Mus. Cont. Biol. GeoL, (42): 1-25, figs. 1-9. Department of EPO Biology , University of Colorado, Boulder ; Colorado, 80309-0334 (HMS); Museo de Zoologla, Facultad de Ciencias , UN AM, Apartado Postal 70-399, Mexico, Distrito Federal 04510 Mexico (OFV); and Department of Psychology, University of Colorado, Boulder, Colorado , 80309-0345 (DC). Received: 17 August 1992 Accepted: 20 September 1992 Bulletin of the Maryland Herpetological Society Page 3 Volume 29 Number 1 March 1993 PHYLOGENETIC RELATIONSHIPS AND TAXONOMY OF COLOSTETHUS MANDELORUM (ANURA: DENDROBATIDAE), WITH NOTES ON COLORATION, NATURAL HISTORY, AND DESCRIPTION OF THE TADPOLE Enrique La Marc a In this paper, the Neotropical frog Colostethus mandelorum is redescribed, and new data on measurements, coloration in life and in preservative, and natural history is added to the knowledge on the species. The tadpole of the species is also described for the first time, and a discussion of phylogenetic relationships is provided, En este trabajo se redescribe la ran a neotropical Colostethus mandelorwru y se complementa el conocimiento sobre la especie con datos nuevos sobre medidas morfometricas, coloracion en viday en preservative, e his tori a natural, Tambien se describe por vez primera el renacuajo de la especie, y se ofrece una discusion sobre sus relaciones filogeneticas. Among the thirty- two frogs collected by E. R. Blake during the 1932 Mandel Expedition to northeastern Venezuela, there were two specimens secured in the mountain known as Cerro Turimi quire (or Turumiquire). They were described by Karl P. Schmidt (1932) as a new species, although unfortunately leaving aside a large series of conspecific frogs discovered two years earlier in the same mountain by M. G. Netting. The new form, Phyllohates mandelorum , was differentiated from P. trinitatis Garman, the geographically nearest congener, on the basis of its dorsal and ventral coloration (Schmidt, 1932: 160). The name was apparently used only once in the literature (Gines, 1959) until it was assigned to the genus Prostherapis and considered a subspecies of P. trinitatis by Rivero (1961: 1 60) . Rivero did not give reasons for usage of the trinomen, but from his “remarks” on the species it could be inferred that the decision resulted from comparing the holotype of Phyllobates mandelorum with syntypes of P. triniiatus (presumably those in the Museum of Comparative Zoology at Harvard), and noting that they had a “striking similarity” (a similarity that he expected on the basis of close proximity to the type-locality of P. L trinitatis ; Rivero, 1961: 161). Since then, the trinomen has been used several times in the literature. Although the name Prostherapis mandelorum was used by Dononso-Barros (1966), and Colostethus mandelorum (current generic allocation) by Edwards (1971), neither author gave reasons for returning to the binomen. Working Page 4 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 independently, several authors (Hardy, 1984: 109; La Marca, 1984: 245; Rivero, 1984: 10) discussed the distinctiveness of the species previously suggested by Edwards (1971). In spite of the general agreement about the validity of the species, the systematic relationships of Colostethus mandelorum are not clear. Thus, Rivero (1984: 12) considered that C. mandelorum most probably had derived from an ancestral stock of C. trtnitatis , a position later sustained when he (Rivero, 1990: 23-24) indicated the possibility that the species is “un derivado de las especies acollaradas.” This consideration now it h si anding , he did not assign the species to any of his later (Rivero, 1990) species groups. Examination of the holotype and paratype of Colostethus mandelorum (Schmidt), along with twenty- two additional specimens, made clear that, although C. mandelorum is a distinct species, its relationships are not to be found among the collared frogs with whom they have been repeatedly related (La Marca, 1984: 245, “ms=3”; see below). These results, along with other data on the species so far unpublished, prompted me to write this paper in the hope that it will contribute to the current taxonomic and systematic knowledge on the species. A detailed description of the holotype is provided to complement two short available descriptions (the original by Schmidt, 1932; and that of Rivero, 1961, that largely duplicates the original). The format, methods, and terminology, as well as pertinent Colostethus material examined in association with this study, are found in La Marca (1984). Staging of tadpoles was performed according to the methods of Gosner (1960). Specimens of C. mandelorum examined for this study are : FMNH (Field Museum of Natural History) 17788, male, holotype* 17789, male, paratype, Estado Sucre, Mount Tummiquire, 8000ft.; ANSP (Academy of Natural Sciences of Philadelphia) 25757, female, Estado Sucre, Mount Turumiquire; CM (Carnegie Museum of Natural History) 9132 ~ 9135, 9137, 9140, 9152 - 9154, 9158 - 9165, 9167, 9174, 9176 - 9177, (5 males, 14 females, 3 juveniles), 9172 (tadpoles), Estado Sucre, Eivecia. Colostethus mandelorum (Schmidt) PhyRohates mandelorum Schmidt, 1932: 160. Prostherapis trinitatts mandelorum , Rivero, 1961: 160. Prostherapis mandelorum , Donoso-Barros, 1966: 29. Colostethus mandelorum, Edwards, 1971: 148; Hardy, 1984: 109; La Marca, Bulletin of the Maryland Herpetoiogical Society Page 5 Volume 29 Number 1 March 1993 1984: 245; Rivero, 1984: 10; Frost, 1985: 91; Rivero, 1990: 23; La Marca, 1992a; 27, Description of the holotype (with notes on intraspeeific variation given within parentheses), -Head as long as wide; top of head smooth, interorbital area convex; interorbital distance about 2 times greater than upper eyelid width; canthus rostralis not well defined; nostrils directed laterally, not elevated (slightly elevated in paratype FMNH 17789); nostrils closer to tip of snout than to eye; loreal region almost flat, sloping abruptly to lip; snout subelliptical in dorsal view; tip of snout broadly rounded in dorsal view (almost truncate in several specimens; e.g. Fig. 2); tip of snout rounded in lateral profile; length of eye greater than eye to nostril distance; intemarial distance greater than eye to nostril distance; tympanum about 1/3 the length of eye, somewhat inconspicuous, with only anterior and ventral parts visible; tympanum separated from eye about 2/3 its horizontal length; no supratympanic fold; a single tubercle at rictus, no post-rictal tubercles; tongue piriform, longer than wide, with posterior end deeply notched (entire in paratype FMNH 17789); posterior 1/3 of tongue not adherent to floor of mouth; choanae rounded, not concealed by palatal shelf of maxillary arch (partially concealed in most female samples); maxilla and premaxilla toothed, teeth pedicelate and short. Dorsum smooth (some specimens bear scattered small tubercles, more conspicuous on lower back); flanks smooth (tuberculate in several specimens); throat, chest and venter smooth; brachium tuberculate, antebrachium smooth; no ulnar fold or row of tubercles; palmar tubercle single, subtriangular (broadly rounded in paratype FMNH 17789) in outline, about 4 times size of thenar; thenar tubercle elongated, about 1.5 times longer than wide, elevated; no supernumerary tubercles; subarticular tubercles moderate-sized, flattened, ovalate; small pads on fingers; largest pad on third finger, almost covering the tympanum when placed on it; pads wider than long; pad on third finger about 1.6 times wider than adjacent phalanx; second and third finger inconspicuously keeled (Fig. 1), first finger shorter than second; third finger not swollen. Cloaca! opening slightly above midlevel of thighs, directed posteroventrally; veiy short cloaca! fold with scalloped free margin; thighs, shanks and tarsi tuberculate above, smooth below; tarsal fold well-defined, not ending in tubercle, extending from the base of the inner metatarsal tubercle obliquely across ventral side of tarsus to the median line; tarsal fold connected with outer lateral flap of inner toe: outer metatarsal tubercle rounded, subconical in lateral profile; inner metatarsal tubercle oval, about twice longer than wide, and about twice as large as the outer metatarsal tubercle; no supernumerary tubercles; subarticular tubercles small, rounded Page 6 Bulletin of the Maryland Herpetoiogical Society Volume 29 Number 1 March 1993 to oval, flattened; toes moderately webbed; web of the holotype matches the low values of the formula for the series studied, which follows: I (1.0- 1.5)- (0.5- 1.0) II (1.0-1. 5H0.5-1.0) III (1.0-1.5M1.G-1.5) IV (0.5-1.0)-(1.0-2.0) V [higher values correspond mostly to Elvecia specimens, although at least one specimen from the type locality, ANSP 25757, possesses a web development similar to the Elvecia sample); toes with flap-like lateral fringes; a lateral keel running from near the outer metatarsal tubercle (or near the mid metatarsus in most specimens) to base of fifth toe, where it meets the outer lateral fringe on that toe; pads slightly wider than long; largest pad on fourth toe, slightly larger than pad on third finger and about 1 .7 times wider than adjacent phalanx (Fig. 1); when thighs are held at right angles to body axis, heels slightly overlap as in other males (in females, either they do not overlap or just touch each other) ; heels reaching to posterior comer of eye when legs are adpressed forward. Measurements of holotype (in mm, abbreviations as in Table 1). -SVL 25.8, IN 3.3, HW 9.2, TL 1 1.5, EN 2.0, HAND 7.6, FOOT 1 1.5, EYE 3.5, testes 3.0. Measurements and proportions for other Colostethus mandelorum (including the holotype) are given in Table 1. Coloration in preservative. (Fig. 2) Dorsum pale cream with dark brown sinuous paravertebral markings (those of the holotype can be best described as an inverted triangle encompassing both upper eyelids, with base toward anterior part, connected with an irregular diamond-shaped marking in shoulders having pale spots inside, and continuing on posterior part with irregular brown markings with ill-defined borders. Brown markings stopping at level of inguinal region, where two dorsal pale bands of background coalesce to form a narrow band on the end of urostyle); dark brown dorsolateral stripe extending from anterior border of eye to inguinal region (some specimens, including the holotype, have the top of anterior part of head pale cream with a brown spot just in front of an imaginary line connecting anterior ends of upper eyelids) ; specimens from Elvecia, agreeing largely in morphology with the holotype, have a larger amount of variation in their pattern of dorsal spots than do topotypes (Juvenile CM 9140, from Elvecia, however, have similar dorsal pattern to that of the holotype); a pale stripe may run (interrupted or not) from insertion of one upper arm to insertion of the other, surrounding the head; darkcanthal stripe surrounding snout and encompassing nostrils; lower lip dusted with brown; dark stripe running on upper lip, sometimes presenting a medial bump on loreal region; tympanum paler than surroundings; a dark longitudinal band on anterior part of upper arm; discrete pale inguinal stripe (occasionally only a series on non-connected pale spots) usually running from inguinal region to point where elbow reaches when adpressed backwards on flank; borders of anal opening pale, sometimes with small pale spots below; dark brown longitudinal Bulletin of the Maryland Herpetologicai Society Paae 7 Volume 29 Number 1 March 1993 stripe on anterior part of thigh; diffuse cross-barrings on thighs, shanks and tarsi (cross-bar markings are more conspicuous in juveniles); concealed parts of tarsi dark, with a pale tarsal fold; adult females with marbled throat, chest and upper venter, and pale cream lower venter; adult males with brownish ventral surfaces where dark brown specks are less conspicuous than, and pale spots smaller than in females; two brown spots on throat, below chin; collar absent; palms and soles darker than other ventral surfaces. Coloration in life.- Schmidt (1932; 160) reported a greenish silvery dorsum and top of the head for the holotype. Other notes on coloration, used for the following color description, are found in Netting’s field notes of January 1930. In these notes, several dorsal color patterns for individuals of C. mandelorum were reported: olive green, bronze, greenish, dark greenish gray, and bright greenish bronze. Flanks were described as greenish, gray green or olive greenish; the inguinal stripe as pale yellow; throat and chest as yellowish, gray, yellow white or pale bluish white; chin orange or pinkish ; markings on chin as bluish white; markings on throat and venter were noted as brown, bluish white, red or reddish brown; venter as pale green, greenish yellow, bright yellow, yellow orange, pale yellow or greenish; upper legs were indicated as brown, and the under legs as pale brown, dull orange, light green or orange; under arms are more orange than yellow; a touch of yellowish at leg insertions, and toe pads were gray. Natural history .- Almost nothing is known on the natural history of Colostethus mandelorum , although a few aspects can be drawn from the available field notes and the localities where they have been taken. Thus, from one of the places (Elvecia, Edo. Sucre) is known that the species occurred in small pools along mountain streams, where they dived and rested on the bottom when they were frightened (Netting, field notes January 1 930) . From the sample available for study, it can be inferred that males may reach sexual maturity at a size of at least 22.6 mm, and females at 24.0 mm, judged from the enlarged testes and vocal slits, and deeply convoluted oviducts and/or mature ova, respectively. Stomachal contents revealed the presence of the following prey items: ants (adult male CM 9135) , Coleoptera (adult females CM 9137 and 9159), and both ants and Coleoptera (adult female CM 9132). Some ecological aspects of the habitat of Colostethus mandelorum may be extrapolated from the places where the species has been collected. Those places lie within the “Macizo del Turimiquire”, an erogenic feature holding at least four life zones. These, in the classification of Ewel et al. (1976) are: premontane humid forest (“bosque humedo premontano”, bh- P), premontane very humid forest (“bosque rriuy humedo premontano”. Page 8 Bulletin of the Maryland Herpetologicai Society Volume 29 Number 1 March 1993 bmh-P), lower montane very humid forest (“bosque muy humedo montano bajo”, brnh-MB) , and lower montane humid forest (“bosque humedo montano bajo”, bh-MB). The mean annual temperatures of these environments range from 1 1 to 24° C, with a mean annual precipitation of 1000 to 4000 mm, and an altitudinal range covering from 500 to 2600 m. Our only datum of altitude for the species is “8000 ft.” (about 2440 m), corresponding to the type specimen. Only two of the mentioned life zones match this elevation, bmh- MB and bh-MB. These ecosystems are immediately above the premontane forests, separated from them by the “critical” temperature (Bwel et al., 1976) of 18° C, a value that seems to be of paramount importance for different species. Thus, for example, coffee plantations grow best in premontane forests, being almost absent in lower montane ones. On the contrary, autochthonous conifers ( Podocarpus spp.) and introduced grasses (like the “Kikuyo” grass, Pennisetum clcmdestiruim) grow best in lower montane forest. This “critical” temperature appears to affect frogs in a similar manner; species of Martnophryne prosper in premontane forests (La Marca, 1992b), while those of the Coiostethus alhoguttatus group are abundant in lower montane forests. In this regard, Coiostethus mandelorurn may occupy habitats similar to those inhabited by the species of the C. alhoguttatus group, while its purported relative, Coiostethus ( ^Mannophryne ) trinitatis , lives in premontane forest; that is, the “critical” temperature of 18° C may be separating the distributional ranges of both species. This hypothesis needs to be tested when more material and data become available. For the moment, considering the evidence, it seems safe to state that Mannophryne inhabit places with a mean annual temperature between 18 and 24° C, while species in the C. alhoguttatus group, as well as Coiostethus mandelorurn, live in places with temperatures between 18 and 11° C. In both cases, a minimum of 1000 mm of mean annual precipitation seems to be germane. These environmental parameters may constitute useful cues of prospective places where to look for these frogs. Tadpole - Nineteen larvae (CM 9172) collected at Elvecia, Estado Sucre, are referred to Coiostethus mandeloram This allocation is tentative because no tadpoles carried on dorsa were available for examination. No other dendrobatid frog is known to occur sympatrically with C. mandelorurn , although Mannophryne trinitatis occurs at lower elevations than C. mandelorurn on Cerro Turimiquire. The tadpoles from Elvecia differ from those of Coiostethus (= Mannophryne) trinitatis reported by Kenny (1969) in having the first upper row of denticles completely keratinized, with equal¬ sized denticles. Description of tadpoles - Body oval in dorsal view, depressed (wider than deep), deepest and widest at about two thirds length of body (from tip of snout to body-tail juncture); tip of snout broadly rounded; lateral line Bulletin of the Maryland Herpetologlcal Society Page 9 Volume 29 Number 1 March 1993 organs appealing as pale dots arising near the mouth, crossing mid-loreal region to surround posterior part of eye and return again to tip of snout, panning parallel to an imaginary line connecting eye and nostril. Another line of pale dots originating posteriorly to eye, bifurcating into dorsolateral and vertical lines, the latter splits into two branches (just above and anteriad to spiracle, on left side), one directed vertically, surrounding base of head, and another running oblique to middle of caudal musculature; nostrils dorsolateral, directed anterolaterally, rounded with pale borders, slightly closer to eye than to tip of snout; intemarial distance wider than interorbital distance; chondrocranial elements not visible through skin of head; spiracle sinistral, forming a tube attached to body wall; spiracular opening directed posterodorsally at about midlength of body and at about 40% distance from bottom to top of body; cloacal tube short, with dextral opening; dorsal fin arising at body-tail juncture or anteriorly; caudal fins about equal in depth; fins shallowe r th an cau dal musculature at midtail length ; caudal mu sc ulature moderately strong, tapering gradually to end just short of tip of tail; tip of tail rounded (Fig. 3, top). Mouth relatively small, situated ventrally at tip of snout, directed anteroventrally; mouth with moderately developed lateral folds; a single row of alternating marginal papillae, except for anterior diastema on upper lip; single row of alternating papillae at comers of lower lip; beaks strong, well keratinized only near cutting edge, bases not keratinized; lower beak with stronger dentition than upper beak; blunt serrations on both beaks; upper beak with narrow lateral extensions; lower beak broadly V-shaped; two upper and three lower rows of denticles; upper rows of same length, arched outward medially; first upper row complete, second upper row widely separated medially; lower rows complete, equal-sized, about equal in length to upper rows; denticles on third row smaller and less keratinized than those on other rows (Fig. 3, bottom). Tadpole coloration - In preservative, dorsum, flanks and venter uniformly dark brown; caudal musculature paler than body; caudal fins cream, dusted brownish, with large dark brown flecks. Coloration in life unknown. Measurements of tadpoles (in mm ± standard deviation).- Stage 25 (N=8) , body length = 8.84 ± 0. 5; total length = 20.9 ± 1 . 1 . Stage 26 (N~5) ; body length = 9.84 ± 0.4; total length = 23.9 ±2.1. Stage 27 (N=6); body length = 11.75 ± 0.5; total length = 29.9 ± 1.5. Discussion - Colostethus mandelonun has usually been associated with a collared frog ( Mannophryne trinitatts) owing to aspects of the ventral Page 10 Bulletin of the Maryland Herpetologicai Society Volume 29 Number 1 March 1993 and dorsal coloration already pointed out by Schmidt (1932: 160). Throat and chest of male hole type and paratype are dark, covered by melanophores, although not as black as in sexually matured male collared frogs (Mannophryne). I regard this condition as different from that exhibited by collared frogs, and therefore I consider Colostethus mandelomm as lacking a”true” collar. The absence of a collar in C. mandelorum is a non- informative character, but the combination of patterns of dorsal color, dark dorsolateral bands, and ventral and gular melanophores may indicate that the species is not closely related to any of the species of Mannophryne. The gular pattern of melanophores in C. mandelorum reminds the condition seen in members of the C. alboguttatus group. This group, designated by La Marca (1985), is diagnosed based on a dentition characterized by the presence of long maxillary and pre-maxillary teeth (“Fang- like” teeth of La Marca, 1984: 245, “ms. 1”). The same character is also exhibited by the monotipic Aromobates Myers et al. ( 199 1). Although its relationships to the Colostethus alboguttatus group can not be ascertained at present, Aromobates noctumus is most similar to some species in this assemblage (like, for e.g., Colostethus leopardalis) . Although Aromobates has a smell that is not present in members of the C. alboguttatus group (probably an apomorphic character), I consider this species to be more closely related to this group. Further studies may even reveal that them all are congeneric. Colostethus mandelorum lacks long teeth, making proper allocation difficult. Characters of C. mandelorum resembling those present in members of the C. alboguttatus group may be plesiomorphic and the species may be either completely unrelated, or a relatively primitive member of that assemblage. Colostethus mandelorum is readily distinguished from species within Mannophryne and within the C. alboguttatus group by the absence of a dark collar, and by the presence of small teeth, respectively. The combination of dorsal pattern of coloration, dark dorsolateral stripes and distribution of melanophores on throat and venter resembles that seen in some members of the C. alboguttatus group but, without knowing the degree of apomorphy of these characters, allocation of C. mandelorum to this group would be unfounded. The presence of lateral fringes along the second and third fingers (Fig. 1) is shared, among other Venezuelan Andean and coastal range Colostethus, only with C. dunni (Rivero) and C. leopardalis Rivero. Colostethus dunni possesses an extensive foot web and different dorsal pattern, and neither C. dunni nor C. leopardalis have dorsolateral stripes. Furthermore, the upper beak in the Elvecia tadpoles has a median convexity resembling that present in some members of the C. alboguttatus group (e.g. Colostethus mayorgau see La Marca and Mijares, 1988: 49). Other traits such as the position of the spiracle, and the type of serration on beaks are also more similar to those shown by members of the C. alboguttatus group. Lack of Bulletin of the Maryland Herpetologica! Society Page 1 1 Volume 29 Number 1 March 1993 knowledge on the distribution of these features among dendrobatids precludes inferences about its importance in determining phylogenetic relationships. Acknowledgments i am grateful to Drs. C. J. McCoy (CM) and T. Uzzell (ANSP) for loan of herpetological material under their care. Dr. McCoy reviewed an earlier version of the manuscript. Dr. Jesus Molinari and Mr. Abraham Mijares provided helpful comments on the final draft. Dr. M. Graham Netting’s generously granted permission to quote from his field notes. Page 12 Bulletin of the Maryland Herpetological Society Bulletin of the Maryland Herpetological Society Page 13 Volume 29 Number 1 March 1993 Fig. 1 Left: Hand of Colostethus mandelorum, CM 9162. Right: foot of C. mandelorurru CM 9135. Scale equals 5 mm. Page 14 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Bulletin of the Maryland Herpetological Society Page 15 Fig. 2. Dorsal and ventral view of Colostethus mandelorum. Left * CM 9174, female, SVL 27.9 mm. Right = CM 9176, female, SVL 24.0 mm. Volume 29 Number 1 March 1993 Page 16 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Literature Cited Donoso-Barros, R i960. Dos nuevos Gonatodes de Venezuela. Museo Nacional de Historia Natural, Santiago de Chile, Publicacion Ocasional 11:1-32. Edwards, S. R 1971. Taxonomic notes on South American Colostethus with descriptions of two new species (Amphibia, Dendrobatidae). Proc. Biol. Soc. Washington 84(18): 147-162. Ewel, J. J., A. Madriz, and J. A. Tosi, Jr. 1976. Zonas de Vida de Venezuela. Memoria explicativa sobre el Mapa Ecologico. 2nd. Ed. M.A.C., FONAIAP, Caracas. 270 p + 1 map. Frost, D. R (Ed.). 1985. Amphibian species of the world. A taxonomic and geographical reference. Allen Press, Inc., and The Association of Systematic Collections, Lawrence, Kansas. 732 p. Gines, Hno. 1959. Familias y generos de Anfibios -Amphibia- de Venezuela. Memoria Sociedad de Ciencias Naturales La Salle 19(53) :84- 146. Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica 16:183-190. Hardy, Jr., J. D. 1984. Systematic status of the South American frog “ Phyliohates mandelorum " (Amphibia, Dendrobatidae). Bull. Maryland Herp. Soc., 20(3): 109- 111. Kenny, J. S. 1 969. The Amphibia of Trinidad. Studies on the fauna of Curacao and other Caribbean islands 108. The Hague. 78 pp + xv pis. Bulletin of the Maryland Herpetological Society Page 17 Volume 29 Number 1 March 1993 La Marca, E. 1984. A taxonomic and systematic revision of the frogs of the Colostethus collaris Group (Anura: Leptodactylidae: Dendrobatinae). Unpubl. Master’s Thesis. Univ. Nebraska. 256 p. 1985. A new species of Colostethus (Anura: Dendrobatidae) from the Cordillera de Merida, Northern Andes, South America. Occas. Pap. Mus. Zool. Univ. Michigan 710:1-10. 1992a. Catalogo taxonomico, Biogeografico y Bibliografico de las ranas de Venezuela. Cuademos Geograficos 9, Fac. Cienc. Forest. Univ. Los Andes, Merida. 1992b. Geografia de las ranas andinas de Venezuela. Geographica de Merida l(l):25-30. _ _ and A. Mijares U. 1 988. Description of the tadpole of Colostethus mayorgai (Anura: Dendrobatidae) with preliminary data on the reproductive biology of the species. Bull. Maryland Herp. Soc., 24(3):47- 57. Myers, C. W., A. Paolillo, and J. W. Daly. 1991. Discovery of a defensively malodorous and nocturnal frog in the family Dendrobatidae: phylogenetic significance of a new genus and species from the Venezuelan Andes. Amer. Mus. Novit., 3002:1-33. Rivero, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126(1): 1- 207. 1984 (“1982”). Sobre el Colostethus mandelorum (Schmidt) y el Colostethus inflexus Rivero (Amphibia, Dendrobatidae). Memoria Soc. Cienc. Nat. La Salle 42(118):9-16. 1990 (“1988”). Sobre las relaciones de las especies del genero Colostethus (Amphibia; Dendrobatidae). Memoria Soc. Cienc. Nat. La Salle 58(129):3-32. Page 18 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Schmidt, K. P. 1932. Reptiles and Amphibians of the Mandel Venezuelan Expedition. Field Mus. Nat. Hist. Zool., Series Pub., 309, 18(7): 159-163. Institute de Geograjla, Universidad de Los Andes . Apartado Postal 116 . Merida 5 10 LA. Venezuela. Received: 13 October 1992 Accepted: 10 November 1992 Bulletin of the Maryland Herpetoiogicai Society Page 19 Volume 29 Number 1 March 1993 VARIATION IN TWO SPECIES (G. BICOLOR, G. DUELLMANI) OF MEXICAN EARTH SNAKES ( GEOPHIS) Hobart M. Smith and Oscar Flores Villela Geophis bicolor is recorded from the state of Mexico, and for the first time from the state of Michoacan, with variations that leave in question the separability of that taxon from G. chalybeus. The second and third males of G. duellmani are recorded with their variations. Among the specimens of Geophis in the Museo de Zoologia of the Facultad de Ciencias (MZFC) of the Universidad Nacional Autonoma de Mexico are four specimens that extend knowledge of the distribution and variation of their two species. Geophis bicolor Gunther Two males are present, one from near the central southern border of the state of Mexico, municipality of Tenancingo, Salto de Agua near Santa Ana (MZFC 41), the other (MZFC 835) from the state of Michoacan, municipality of Zitacuaro, 0.5 km SSE San Francisco de Coatepec. The former was previously reported by Lara and Flores (1980: 371), although without description, and the latter is the first known from Michoacan. Respectively, the ventrals are 1 56, 158; caudals 51,47; total ventral- caudals, 207, 205; scale rows 17, 17; s-v 189, 103 mm; tail 52, 22 mm; total length 241, 125 mm. In MZFC 41, the postoculars are 1-1, an anterior temporal is present on one side (separating parietal from 5th supralabial), and the two prefrontals are half fused. MZFC 835 has a tiny, lower postocular on one side, and the other side is damaged, the number of postoculars indeterminate, although if a lower postocular is present on that side it is tiny, like the other, because a very large, single postocular is evident as on the intact side above the position where a tiny lower postocular may have been before the head was damaged. The dorsum in both specimens is uniform dark brown except for light centers on the lower 2-4 rows of dorsals. The venter is unpigmented, and in MZFC 41 the subcaudal surface is irregularly pigmented. Page 20 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Downs (1967: 56) noted one other specimen with an anterior temporal on one side, but no other with any fusion of the prefrontals, and none with only 1-1 postoculars or with such a tiny lower postocular. Indeed, presence of two postoculars is one of the few distinctions of G. bicolor from G. chcdybeus , as determined by Downs (1967: 55-59). Data are available only for three of the latter taxon, however, all from Mirador, Veracruz, and for 16 G. bicolor (eight in Downs, 1967, and eight in Dixon, 1968), all from either Jalisco or “neighborhood of Mexico City,” the latter of which we regard as erroneous (although Downs, 1967: 57 argued for its validity). All previously recorded G. bicolor have two postoculars except on one side of two, with one (Dixon, 1968: 453), and all have extensive ventral pigmentation except for those from “neighborhood of Mexico City.” The present two specimens, from localities intermediate between Jalisco (G. bicolor) and Veracruz (G. chalybeus ), resemble the latter species more than the former in having the venter essentially immaculate (as do those supposedly from near Mexico city), and in having 1-1 postoculars or only a tiny lower post ocular in addition to a large upper one. The only apparent distinction remaining between G. bicolor and G. chalybeus , as interpreted by Downs (1967), is the lower ventral count in females (the only sex yet recorded) of the latter ( 1 54- 1 55 vs 1 6 1 - 1 68) and its generally lower ventral plus caudal count (192-196 vs. 195-209). Other apparent differences evident in comparing Downs’ two descriptions (1967: 55-59) are not borne out by Dixon’s (1968) detailed description of Jalisco specimens, or by our own material. It is therefore distinctly possible that G. bicolor Gunther, 1868, is a synonym or geographic race of G. chalybeus (Wagler, 1830), inasmuch as their known ranges are allopatric (whether in reality significantly dichopatric, or essentially parapatric, is unknown) and only ventral counts now appear to differ. Indeed, on the contrary, postocular counts and ventral pigmentation suggest that the populations inhabiting the environs of Jalisco may well be taxonomically distinct from either of the other two, at least at subspecific rank. Clearly more material, particularly of G. chalybeus auct., is essential to clarify the taxonomy of this complex. Until that time we maintain the taxonomic status quo, because adoption of any alternative would be essentially an exchange of one uncertainty for another, with its accompanying nomenclatural vacillation and confusion. Bulletin of the Maryland Herpetological Society Page 21 Volume 29 Number 1 March 1993 Geophis duellmani Smith and Holland Two adults (MZFC 5081), both DOR males, are from La Esperanza, 1700 m, 80.5 km S Tuxtepec, Oaxaca, less than a km from the type locality, 2 mi S Vista Hermosa (apx. 3 km N La Esperanza) . Only four other specimens have been reported in the literature (Campbell et al., 1983), all from 2.8 to 6.6 km S Vista Hermosa, but including only one male. The present two accordingly extend the known range of certain characters of males: ventrals 126.5, 129 (from 133); subcaudals 40, 43 (from 43); dark saddles 16-18 on the two sides of one, 21-22 on the other (from 18-21 in both sexes). Other characters conform with previous descriptions. Total lengths are 214 and 230 mm, tall 42 and 47 mm respectively. Acknowledgments We are grateful for the support to OFV from DGAPA-UNAM project no. IN201789, and to HMS for research facilities provided through the generosity of Dr. William M. Lewis, Chairman of the Department of Environmental, Population and Organismic Biology, and Dr. Shi-Kuei Wu, Curator of Zoology in the Museum, both at the University of Colorado. yiejat.uie.£it^d Campbell, Jonathan A., Linda S. Ford and John P. Karges. 1 983 . Resurrection of Geophis anocularis Dunn with comments on its relationships and natural history, Trans. Kansas Acad. Sci., 86(1): 38-47, figs. 1-2. Dixon, James R. 1968. Notes on the snake genus Geophis , from Nevado de Colima. Jalisco. SW Naturalist, 13(4): 452-454. Downs, Floyd L. 1967. Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Misc. Publ. Mus. Zool. Univ. Michigan, (131): i-iv, 1-193, figs. 1-23. Page 22 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Lara Gongora, Guillermo and Oscar Flores Villela. 1980. Primer catalogo de una coleccion herpetologica depositada en la Facultad de Ciencias. Mem. II Congreso Nac. Zool. 1978, 2: 366-374. Department of Environmental, Population and Organismic Biology, University of Colorado, Boulder, Colorado 80309 0334 U.SJl. (HMS); Museo de Zoologta, Facultad de Ciencias, Universidad Nacioncd Auto noma de Mexico, Apartado Postal 70 399, Mexico, Distrito Federal, 04510 Mexico (OFV). Received: 16 October 1992 Accepted: 7 December 1992 Bulletin of the Maryland Herpetological Society Page 23 Volume 29 Number 1 March 1993 OBSERVATIONS ON THE SENSITIVITY TO HIGH TEMPERATURES IN TWO LIZARD SPECIES (AMEIVA UNDULATA AND SCELOPORUS HORRIDUS) FROM ZACATEPEC, MORELOS, MEXICO Julio A. Lemos- Espinal, Royce E. Ballinger and Juan M. Javelly-Gurria Studies on lizard thermal physiology have increased since the early work of Cowles and Bogert (1944). However, there has been relatively little work on thermal characteristics of Mexican lizard species beyond an initial study by Bogert (1949) on the genus Sceloporus. In his classic study, Brattstrom (1965) reported the eccritic body temperatures and upper thermal limits for some Mexican lizard species. More recently Ballinger et al. (in manuscript) documented active body temperature and heat tolerance for Xenosaurus grandis located at Cuautlapan, Veracruz, Mexico, and Lernos- Espinal ( 1992) compared active body temperatures and heat tolerance of two populations of the high altitude lizard Sceloporus grammicus on the Iztaccihuatl Volcano, Puebla, Mexico. Here we report sensitivity to high temperatures in two lizard species ( Ameiva undulata and Sceloporus horridus ) from the vicinity of the city of Zacatepec, Morelos. In general, teiids are active at higher temperatures and have higher thermal sensitivities than iguanids (Brattstrom 1 965) . However, no studies have specifically examined sensitivity to high temperatures in these species at a single location and time, where exposure to prevailing thermal conditions that affect physiological acclimation would be the same. Between June 15-20, 1992, twenty adult Ameiva undulata (average snout-vent length 109. 1 mm ± 2. 10 SE, and body mass 37.17 g ± 2.44 SE) and fifteen Sceloporus horridus (average snout-vent length 95.33 mm ±1 .95 SE, and body mass 35.53 g ± 1.95 SE) were collected at 910 m elevation in the Camp de Investigaciones Forestales y Agropecuarias de Zacatepec, Morelos (CIFAZM) . Individual Ameiva undulatawere captured by hand. They are commonly observed on the ground close to artificial water canals that surround the facilities of CIFAZM. Individual Sceloporus horridus are commonly observed on walls of CIFAZM and were captured by hand or noose. Average body temperatures were significantly higher in Ameiva undulata than in Sceloporus horridus (Table 1). Since all lizards we collected required considerable effort, the body temperatures we obtained reflect high metabolic activity and do not represent equilibrium eccritic body temperatures. Lizards were placed in lab conditions at 25° C and were tested for Page 24 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 sensitivity to high temperatures within 12 hrs of collection. Lizards were heated continuously with a lamp (250 W) suspended 20 cm above the lizard. The lizard was tethered on a surface covered with aluminum foil to avoid substrate heating. We recorded the body temperature (cloacal temperature) with a quick reading Schultheis thermometer at four sequential behaviors (Hertz etal. 1979): (1) Experimental voluntary maximum (EVM) was the body temperature at which an individual lizard first attempted to escape the heat source (Ruibal 1961), (2) Continuous escape (CE) was an escape reaction period characterized by continuous running activity (Ballinger and Schrank 1970), (3) Panting threshold (PT) was recorded as the temperature when the lizard gaped and breathed deeply (Heatwole et a!. 1973), and (4) The critical thermal maximum (CTM) was recorded as the temperature at which locomotor activity became so disorganized that the animal could not escape conditions that would promptly lead to its death as evidenced by paralysis and loss of a righting response (Pough and Cans 1982). After CTMs were noted, lizards were quickly removed from the heat source and allowed to recover. Full recovery occurred within minutes in all lizards tested. Sensitivity to high temperatures between species was compared using analyses of variance (AN OVA) . There were significant differences in the tolerance to high temperatures in all behaviors (Table 1). Ameiva undulata tolerated higher temperatures than Scelopoms horridus. These results are consistent with other studies on teiids that have reported CTMs as high as 46.2° C in Ameiva ameiva , 45.1° C in Ameiva quadrilmeata (Brattstrom 1965), and 51.0° C in Cnemidophorus sexlineatus (Paulissen 1988). On the other hand, CTMs for Scelopoms lizards range from 41.17 for the high altitude lizard Scelopoms grommicus (Lemos-Espinal 1992) to 46.8° C in Scelopoms occidentalis (Brattstrom 1965). Ecologically limiting temperatures, as represented by the CTM, are clearly lower in Scelopoms than Ameiva. Data reported here on species from the same locality and time suggest that the differences in thermal sensitivity have a genetic or phylogenetic basis since both species have access to the same environmental conditions. However, Hertz (1979) found that thermal sensitivities among Anolis species were predicted better by ecological distributions than phytogeny. Whether the differences we observed are a result mediated by behavioral choices of activity times and thermal preferences or by physiological differences in metabolic pathways remains to be examined. Bulletin of the Maryland Herpetoiogical Society Page 25 Volume 29 Number 1 March 1993 Acknowledgments Worker Sr, Carmelo of the Camp de Investigaciones Forestales y Agropeeuarias de Zaeatepec, Morelos, kindly provided field assistance. We thank Geoff Smith for comments on the manuscript. Field work was supported by Institute Nacional de Investigaciones Forestales y Agropeeuarias (INIFAP), and Consejo Nacional de Ciencia y Tecnologia (CONACyT) both from Mexico. Page 26 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Table 1. Indices of heat tolerance in Ameiva undulata and Sceloporus horridus from Zacatepec, Morelos, Mexico. Mean temperatures (Celsius ± 2 se) are given for experimental voluntary maximum (EVM), continuous escape (CE), panting threshold (PT), and critical thermal maximum (CTM). FIELD TEMPERATURES1 Species A. undulata S. horridus Fl,33 BODY 39.60 ±6.34 35.74 ±0.41 51.86*** AIR 29.68 ±0.61 27.86 ±0.85 3.17NS SUBSTRATE 33.81 ±0.82 30.07 ± 0.95 8.83** SENSITIVITY TO HIGH TEMPERATURES EVM 39.34 ±0.58 36.38 ±0.28 66.71*** CE 41.25 ±0.39 38.77 ±0.17 106.97*** PT 42.50 ±0.32 40.12 ±0.22 129.25*** CTM 45.39 ±0.37 43.22 ±0.37 64.46*** * Temperatures represent high metabolic activity (continuous running). NS * Not significant* ** P < 0.01; *** P < 0.001. Bulletin of the Maryland Herpetoiogical Society Page 27 VoSume 29 Number 1 March 1993 Literature Cited Ballinger, R. E., and G. D. Schrank. 1970. Acclimation rate and variability of the critical thermal maximum in the lizard Phry nosoma comutum. Physiological Zoology 43: 19-22. Ballinger, R. E., J. A. Lemos-Espinal, 3. Sanoja-Sarabia, and R. Coady. Ecology of the lizard, Xenosaurus grandis, in the Tropical Deciduous forest of Cuautlapan, Veracruz, Mexico, (in manuscript). Bogert, C. M. 1 949 . Thermoregulation and eccritic body temperatures in Mexican lizards of the genus Sceloporus. Ann. Inst. Biol. Mex. 20:415- 426. Brattstrom, B. M. 1965. Body temperatures of reptiles. Am. Midi. Nat. 73:376-422. Cowles, R. B., and C. M. Bogert. 1944. A preliminary study of thermal requirements of desert reptiles. Bull. Am. Mus. Nat. Hist. 83:261-269. Heatwole, H., B. T. Firth, and G. J. Webb. 1973. Panting thresholds of lizards I. Some methodological and internal influences on the panting thresholds of an agamid, Amphibolorus muricatus. Comp. Biochem. Physiol. 46A.:799- 826. Hertz, P. E. 1979. Sensitivity to high temperatures in three West Indian Grass anoles (Sauna: Iguanidae), with a review of heat sensitivity in the genus Anolis. Comp. Biochem. Physiol. 63A;2 17-222. _ _ A. Arce -Hernandez, J. Ramirez- Vazquez, W. Tirado-Rivera, and L. Vazquez-Vives. 1979. Geographic variation of heat sensitivity and water loss rates in the tropical lizard Anolis gundlachl Comp. Biochem. Physiol. 62A: 947-953. Page 28 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Lemos-Espinal,. J. A i 992 . Ecology and comparative demography of the lizard Scetoporus grammicus: Life history of an altitudinal generalist on the eastern slope of the Iztaccihuatl Volcano, Puebla, Mexico. Unpubl. Ph.D. Dissertation, University of Nebraska, Lincoln. Paulissen, A. M. 1988. Ontogenetic comparison of body t emperature selection and thermal tolerance of Cemidophorus sexUneatus. J. HerpetoL 22:473-476. Pough, F.. H„ and C. Cans. 1982. lire vocabulary of reptilian thermoregulation. In C. Cans and F. H. Pough (eds.)f Biology of the Reptilia, vol. 12. Physiology, pp. 17-23. Academic Press, New York. Ruibal, R 1961. Thermal relations of five species of tropical lizards. Evolution 15:98-111. School of Biological Sciences , University of Nebraska Lincobr Lincoln. Nebraska 68588 A JS A (JAL K and KBB); Centro de Investigaciones Forestales y Agropecuarias de Zacatepec . Morelos , Mexico . Domicilio Coriocido, Apartado Postal # 32780 . Zacatepec , Morelos , Mexico » present address: Centro de Investigaciones Forestales y Agropecuarias del Distrito Federal Ac. Progreso #5, Viveros de Coyoacarc Mexico 0411 (JMJ G). Received: 3 November 1992 Accepted: 7 December 1992 Bulletin of the Maryland HerpetoJogical Society Page 29 Volume 29 Number 1 March 1993 THE RANGE OF THE SO-CALLED RELICTUAL INTERGRADES BETWEEN THE LIZARDS SCELOPORUS UNDULATUS GARMANI AND S. {/. ERYTHROCHEILUS Hobart M. Smith, David Chiszar, Emmett Evanoff and Jeffry B. Mitton The so-called intergrades occur only on slabby, moderately well indurated sandstone ledges associated with loose, highly sandy soils, exposed over at least a dozen meters of extent, where skunkbushes are abundant, in a range including no more than 5% of 25 square miles. The taxonomic status of the population remains uncertain, although it may be resolved by electrophoretic studies now in progress. Intermediacy between S. il garmani and S. u. erythrocheilus now seems unlikely; they represent different exerges and act as different species at their latitudinal levels. More likely is intermediacy between S. u. garmani and S. il consobrinus, both of the central exerge, and the latter (or its progenitors) having given rise to the former. Ever since the extraordinarily stenotopic and stenochoroic (Gr. steno, restricted; choros, land, country) population of Sceloporus undulatus (Bose and Daudin) was accidentally discovered in June, 1989 (Smith et al., 1991), efforts have continued to establish the range of that population. We have now bracketed its distribution, and here summarize present knowledge. In the original report (Smith et al., 1991), the “intergrades” between Sceloporus undulatus erythrocheilus Maslin and S. u. garmani Boulenger were noted as apparently restricted to Fox Hills Sandstone exposures, shunning the extensive limestone outcrops bordering the sandstone habitat to the east and even inter digitating within some areas. That description is in error, to the extent that no limestone occurs in the area. Two major sandstone formations are exposed in the area: the Miocene Ogallala Formation and the Cretaceous Fox Hills Sandstone. The Ogallala Formation is primarily composed of conglomerate and conglomeritic sandstones that are arkosic (derived from granite) and strongly cemented with calcite. Ogallala rocks weather into veiy large boulders. The Fox Hills is composed of quartzose sandstones of two basic types. One type is fine¬ grained, well indurated from calcite cement, and weathers into slabs. The other type is coarser grained and is indurated to varying hardnessess by iron oxide cement. Iron oxide sandstones weather either into hard blocks or loose Page 30 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 sand. Adjacent to these formations (Fig. i) lie extensive Cretaceous deposits of the younger Laramie Formation, composed of coals, carbonaceous shales, mudstones, and muddy sandstones, and the older Pierre Shale, composed mostly of dark, highly organic shales. Three factors appear to be critical in choice of habitat by lizards: 1) presence of hibemacula, as deep crevices in sandstone outcrops: 2) accessibility of a loose, sandy soil for nesting: and 3) the presence of skunkbush ( Rhus trUobata I for cover. The lizards are most abundant on the calcite and iron-oxide cemented Fox Hills sandstones that weather into siabby, moderately well indurated ledges associated with loose, highly sandy sods. Hard, blocks dark-colored sandstone outcrops and siabby outcrops associated with hard, compact soils are not utilized by the lizards. Absence of the lizards in Ogallala sandstone is probably due to a combination of a paucity of hibemacula (Ogallala sandstones lack suitable crevices) and their association with compact, gravelly soils resistant to nest excavation. Skunkbush abounds in most places on both the Fox Hills and Ogallala sandstones, and does not seem to be a limiting factor. Small, isolated sandstone exposures likewise are not inhabited by S celoporus. The lizards are most abundant where continuous stretches of sandstone exist, of a dozen meters or more. Even there the animals may not occur, if the area is relatively barren. The ideal condition appears to require numerous skunkbushes growing at the edges of sandstone slabs. The lizards appear to thermoregulate by alternating between the sunlight and shade that dapple the rock surface under the bushes. Certainly they wander a bit from the shelter of those bushes over rocks, but they seem not to linger long elsewhere, except for females seeking sand in which to lay their eggs. Where yuccas are the major potential sheltering plant over sandstone, the lizards do not occur. Even where conditions appear to be most favorable, with widespread sandstone exposures and abundant skunkbush, the lizards are difficult to find. They are rarely seen unless they move, hence the most effective approach is to scan sharply the overhang area of bushes over rocks, while carefully prodding or trampling the vegetation. The lizards are thus induced to move and thereby are detected. They are very secretive, usually not exposing themselves, and in addition appear to be scarce. Three or four lizards an hour is a good rate of discovery. The accompanying map (Fig. 1) indicates all of the localities where the "intergr ades" have so far been taken. We did not find them where the circles are placed near the bottom of the map. The southernmost pair of those . circles indicates a very favorable-appearing exposure of sandstone Bulletin of the Maryland Herpetological Society Page 31 Volume 29 Number 1 March 1993 caprock south of I 70; it is likely that the absence of these lizards is due to the nearly complete absence of skunkbush. Yuccas are present in abundance, many bordering sandstone slabs, but apparently the density of their shade and cover is unacceptable; otherwise the habitat seems suitable. The area is overgrazed, and that may account for the rarity of skunkbush. The other pair of circles, north of I 70, represents an extensive exposure seemingly veiy similar to the lizardless exposures south of I 70. It, however, consists almost entirely of a thin layer of fine-grained, strongly indurated, calcareous sandstone, where crevices do exist that seemingly would suffice for overwintering. Skunkbush is also abundant. The habitat appears to be suitable for these Scelopoms , except for the compacted soil produced by the weathered sandstone. Whatever the reason, no Scelopoms are there. The small exposure on the eastern end of the ridge is strongly indurated, iron oxide -cemented sandstone, and free of crevices. Its small extent and absence of crevices make it unsuitable for these lizards. Large, similar exposures of exceptionally hard, blackened sandstone occur farther southwest (8-16 km) near state highway 86 to Kiowa, but are devoid of Scelopoms. Isolated exposures of sandstone with skunkbush were examined 14 air miles west and three miles south of Deer Trail (12 mi NNW Agate), on escarpments bordering West Bijou Creek, without success. Extensive sand deposits in the same general area yielded no evidence of Scelopoms. The extensive exposures of Ogallala sandstone on the east escarpment of Beaver Creek are devoid of Scelopoms. The northern circles mark the areas of exposed, flat, fine-grained, calcareous sandstone where no Scelopoms were found. It is our impression, therefore, that the “intergrades” are restricted to the general area shown in Fig. 1 , where we have found them. To the north, no rock exposures exist. To the east, valleys have no rock exposures, and hills are capped with uninhabited Ogallala sandstone. To the south, no favorable habitat exists in spite of propitious topography and rock outcrops. And to the west, only weakly dissected, mostly flat farmland exists. The “intergrade” population therefore seems to be limited to the north side of the terminus of the Palmer Divide, which separates the South Platte River drainage to the north from the Arkansas River drainage to the south. The range includes hills bordering the headwaters of Rattlesnake Creek, extending to the west slopes of Beaver Creek to the east, and the east slopes of Muddy Creek to the west. In all, the range of the intergrade population probably does not exceed 25 square miles, and within that territory the lizards are confined to a particular type of sandstone exposure that does not comprise more than Page 32 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 at most 5 per cent of the total area. The number of individuals is no more, we estimate, than at most 5000. The population is in jeopardy because of its small size and limited habitat. However, its main threat seems to be overgrazing (which risk is minimized by the rough topography) and a hypothermic climatic change. With a long-term hypsithermal climatic change, its present tenuous existence might improve. Its predators (raptors, snakes, carnivores) are likely to be held at bay to some extent by human interference, which in turn is not likely to increase greatly through the foreseeable future in this rugged ranching land, totally unsuitable for farming. The taxonomic status of the “intergrade” population remains a question. It appears superficially simply like S. u. garmani from which it differs primarily in behavior and habitat, secondarily in development of ventral semeions in both sexes, and to a minor degree in femoral pore and interfemoral scale counts (Smith et ah, 1991). Isozyme electrophoretic comparisons with S, u. garmani and S. u. erythrocheilus , now in progress, may provide useful information in this context. Comparative ethologies are also contemplated. Such studies should narrow the range of possibilities relative to taxonomic status of the “intergrades.” At present those possibilities are: (1) as previously hypothesized, relictual intergrades between S, u. garmani and S. u. erythrocheilus; (2) a peripheral variant of S.u. garmani with no relation to S. u. erythrocheilus; and (3) relictual intermediates, taxonomically distinct or not, between S. u. garmani and S. u. consobrinus Baird and Girard, from which the former was presumably derived, representing an intermediate stage in reduction of its semeions (well developed in the latter taxon). Similarity of the “intergrades” to S. u. erythrocheilus lies primarily in the hypertrophied semeions, as compared with S. u . garmani in which they are poorly developed, and these were the two taxa in which relationships were first sought, since they are the ones adjacent to the “intergrade's” range. However, accentuated semeions are characteristic of members of the S. undulatus group as a whole; in S. u. garmani they are greatly reduced. Hence the “intergrades” might well be interpreted as simply an isolated remnant of an ancestral population, with modest semeions, that elsewhere continued the reductional trend in semeion development, becoming the current S. u. garmani Considerations of the proper nomenclature for the “intergrades” should also take into account the deeply ingrained behavioral differences between the three subspecies groups (exerges) of S. undulatus . The eastern exerge fS. u. hyacinthinus [Green], S. u. undulatus) is scansorial (climbing) in habits, mostly on trees: the western exerge (S. u. elongatus Stejneger, Bulletin of the Maryland Herpetological Society Page 33 Volume 29 Number 1 March 1993 S. u. erythrocheilus, S. u. trisiichus Cope) is also scansorial, on both trees and rocks, but mostly the latter; and the central exerge (S. u. consobrinus , S. u. cowlesi Lowe and Norris, S. u. garmani, S. u. tedbrowni Smith et al.) is terrestrial. In this context, the “intergrades” are terrestrial, not scansorial, in spite of being partial to a rocky habitat. For them, rocks are a part of a terrestrial habitat, to which they are mostly confined; their behavior is definitely not scansorial. In that context, they are clearly a member of the central exerge, and therefore it now seems very unlikely that they represent intermediates between that exerge and the western one, because at the latitudes where S. u. garmant (central exerge) and S. u. erythrocheilus (western exerge) occur, they act as different species, maintaining their extensive differences in ranges coming within at least very few kilometers of each other. They are not yet known to be sympatric, but they probably are. Blending of the two exerges occurs farther south, between S. u. consobrinus (central exerge) and S. u. tristichus (western exerge). Nevertheless, the Great Plains, where the “intergrades” now occur, were forested or wooded during the late Wisconsin glacial epoch (Axelrod, 1985), providing a habitat where some member (presumably S. u. erythrocheilus or its forerunner) of the western exerge may have occurred. As the forest subsequently receded to the east and west, while the Great Plains expanded between, ample opportunity may have existed for reproductive contacts to have occurred between the evolving scansorial and terrestrial exerges, even though at present there is none at least between S. u. garmani and S. u. hyacinthinus on the east, the former and S. u. erythrocheilus on the west. Thus none of the three possibilities, relative to the ancestry and current taxonomic status of the so-called “intergrades”, can be conclusively accepted over the others on the basis of present knowledge. Electrophoretic studies in progress may well restrict the possibilities. Acknowledgments We are much indebted to our field companions and counselors, especially Carl E. Bock, Adam and Dan Chiszar, David Drew, Christine M. Dwyer, H. William Peterson, Matthew S. Rand, and Bruce D. Smith; to William M. Lewis for facilities; and to Shi-Kuei Wu for curating material. Page 34 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 Fig. 1 . Distribution of the isolated population of Scelopoms undulatus near Agate, Elbert Co., Colorado. Map traced from the Bureau of Land Management's 1985 Limon and Last Chance 1:100,000 scale metric topographic maps. Geology after Sharps (1980). Solid dots indicate sites where samples were found, hollow dots sites where habitats might be suitable but where no lizards were found. Bulletin of the Maryland Herpetoiogical Society Page 35 Volume 29 Number 1 March 1993 Literature Cited Axelrod, Daniel L 1985. Rise of the grassland biome, central North America. Bot. Rev., 51: 163-201, figs. 1-4. Sharps, Joseph A. 1980. Geologic map of the Limon 1°X 2° quadrangle, Colorado and Kansas. U. S, Geological Survey Misc. Investigations Map 1- 1250, scale 1 : 250,000: 1 sheet. Smith, Hobart M., Matthew S. Rand, J. David Drew, Bruce D. Smith, David Chiszar and Christine M. Dwyer. 1991. Reiictual intergrades between the Northern prairie lizard (Sceioporus undulatus gcmnani} and the Red-lipped plateau lizard (S. u. erythrocheUus) in Colorado. MW Nat., 72: 1-11, figs. 1-8. Department of Environmental Population and Organismic Biology , University of Colorado, Boulder , Colorado 80309-0334 (HMS, JBM): University of Colorado Museum, ibid., 80309-0218 (DC, EE, HMS); and Department of Psychology , ibid, 80309-0345 (DC). Received: 23 November 1992 Accepted: 7 December 1992 Page 36 Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 £ £ g o 2, a® c >o c ^ « a x* w > TJ © + © © "3 jr ** |jg| © OS 3 £ a -s II 5 HI ® S O K © © “O !« 11 c = *ii 3 ffi t a g o © c S o “ ?s § = § ® O (0 -r o^B 2 « p C 3 5 © S ® > © X c ■» o a o *3 © CB JZ. a, ® © |ll = < ® © o TJ ® £ 8 1 1 1 ! is3* 2 © ° © § TJ © -O ® £ ffi « C*04 | C ffi ™ &t m ® f! E c 2 ^ ® 2 — > © TJ — « O >» © ® © 3 C 3 3 S C © O..S > -c o « Z, Olu®0 « £ ® E o. | 5 g I >* 5 10 « 13 >> ^SS* &ll a SgilSS > a © • g 5 £ « E o|S © «- « © ^ g ~1 o*| f Hip! Ifipi TJ *3 ® O © - „ B ® 5 e o ® cs « £ &£afS 2 o i 5 — © s’ © -8 1 < o a « © 5 S I m ™ >• 3 « !§L2 111 © « 5 p ® . 003 S ^ ® o.-g ss 8 s f Ijl e ; Sf =155 £2* f 8 J s 2 != » « o 3 © © s» §.©•“« ^ ^ ^ c a, ® 5 £8 ® m ® c S jz © ! 1 1- € O 2 « TJ a.% E | a Z 3^ ss o e ~ ® & a ® £ a 1 s 5 O II ® O 2 3 3 •s E g • §• Is it c c =5 £ II © il _a © 3 of 3 © a. c . >N 9 c © c m Q ® p 2 « E © © .o £ # i © 2 c , C ^ Q © t* -SI ‘ > o c» a» cj « « £ . O os _! 03 ® cd^. 3 Si, >, < 8§ §1 Hi B S *** ^ « ssallll «l|ll£ aofi 0*5 g§I“a§5 si .Jigs gsSSeiS^ ** 2** o 2 > s? Page Bulletin of the Maryland Herpetological Society Volume 29 Number 1 March 1993 9\ COTES N A A L II ^ RiSCH !? USEUy WORLD CONGRESS OF HERPETOLOGY Call for nominations for membership of the Executive and International Eerpetological Consort tee . During the Second World Congress of Herpetology, to be held in Adelaide, Australia between Decanber 29, 1993 and January 6, 1994, half of the members of the Executive Ccrrmittee and half of the members of the International Herpetological Garmittee will retire. Membership to both these committees is open to all herpetologists and should be as representative as possible of herpetological subdisciplines and the geographic distribution of herpetologists. Retiring members are eligible for re-election. Nominations to fill the vacancies can be taken now till August 29, 1993. Nominations must be seconded either by any two members of either the Executive or International Herpetological Committees, or, in the case of the International Herpetological Committee, by the governing body of an Affiliated Organisation. Where the number of nominations exceeds the number of vacancies then the election will be by open ballot (during the business meeting) of a format to be determined by the Executive Committee. Present Executive Committee members (with asterisk retiring at SWCH) : R. Avery, UK; W. Bdhme, Germany; * *D.G.Broadley, Zimbabwe; *R.L. Carroll, Canada (Treasurer) ; *G. Casas-Andreu, Mexico; *K.G.Cogger, Australia; *J.C. Daniel, India; *I.S.Darevsky, Russia; W.E.Duellman, USA; T.R.Halliday, UK; *M.S.Hoogmoed, Netherlands (Secretary General); K.Klenmer, Germany; *M.R.K. Lambert, U.K.; J. Lescure, France; J.M. Savage, USA; *P.E.Vanzolini, Brazil; M.H.Wake, USA; Y.L. Werner, Israel; R. Whitaker, India; *E.Zhao, China. Present International Herpetological Committee members: R .A. Acuna Me sen, Costa Rica; P.Alberch, Spain; *J.-L.Amiet, Cameroon; N.Ananjeva, Russia; C.Andr^n, Sweden;* H.Amold, UK; A.M.Ba£z, Argentina; * I. Bar an, Turkey; *W.Be9'b BULLETIN OF THE ^Rarylanb f)erpetolog'ical ©octety DEPARTMENT OF HERPETOLOGY THE NATURAL HISTORY SOCIETY OF MARYLAND, INC. MdHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League 30 JUNE 1993 VOLUMF 90 Ml IMBED O BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY Volume 29 Number 2 June 1993 CONTENTS Sleep-Like Behavior in Young, Captive Red-eared Sliders ( Trachemys scripta elegans) Floyd E. Hayes . . 39 First Record of Amphisbaena dubia Muller (Reptilia: Amphisbaenia) in Argentina A. Alberto Yanosky, James R. Dixon, and Claudia Mercolli .47 A New Species of Knobscale Lizard (Reptilia: Xenosauridae) from Mexico Hobart M. Smith and John B. Iverson . .....51 Observation and Review of the Nesting and Egg-laying of Corytophanes cristatus (Iguanidae). Marco A. Lazcano-Barrero and Eleuterio Gongora-Arones ... 67 The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc. 2643 North Charles Street Baltimore, Maryland 21218 BULLETIN OF THE Volume 29 Number 2 June 1993 The Maryland Herpetological Society Department of Herpetology, Natural History Society of Maryland, Inc. Executive Editor. . .....Herbert S. Harris, Jr. Steering Committee Frank Groves Jerry D. Hardy, Jr. Herbert S. Harris, Jr. Library of Congress Catalog Card Number: 76-93458 Membership Rates Membership in the Maryland Herpetological Society is $16.00 per year. Foreign $20.00/year. Make all checks payable to the Natural History Society of Maryland, Inc. Meetings The third Wednesday of each month, 8:15 p.m. at the Natural History Society of Maryland (except May-August, third Saturday of each month, 8:00 a.rn.). The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 p.m. Volume 29 Number 2 June 1993 SLEEP-LIKE BEHAVIOR IN YOUNG, CAPTIVE RED-EARED SLIDERS (TRACHEMYS SCRIPTA ELEGANS) Floyd E. Hayes Sleep-like behavior was studied in young, captive red-eared sliders ( Trachemys scripta elegans), and compared with an earlier field study on Galapagos tortoises ( Geochelone elephantopus uandenburghi ). The turtles were nearly always “awake9* during the day and “asleep” during the night; when “asleep” they were usually completely submerged. When basking on land during the day, T. scripta usually remained vigilant whereas G. elephantopus usually exhibited sleep-like behavior; differences in the antipredator strategies of these species may explain the differences in vigilance during basking. Both species usually basked with the head and limbs extended, and slept at night with head and limbs retracted; these postures presumably serve a common thermoregulatory function. The electrophysiological presence of sleep has been documented in a variety of turtle species (e.g., Herman et al., 1964; Vasilescu, 1970; Karmanova and Chumosov, 1972; Flanigan, 1974; Flanigan et al., 1974; Ayala-Guerrero, 1987, 1988). But because most previous studies of chelonian sleep focused on the electroencephalographic correlates of specific sleep states, much remains to be learned about the ecological significance of behavioral sleep in chelonians. Hayes et al. (1992) briefly studied sleep behavior in wild Galapagos tortoises (Geochelone elephantopus vandenhwghfl , and found that the tortoises generally slept at midday (apparently while basking) with the head and limbs extended, and slept at night with the head and limbs withdrawn. However, in the absence of electroencephalographic data, they preferred using the term “sleep-like” behavior. As with most species of turtles, previous reports of sleep-like behavior in the slider turtle [Trachemys scripta} are anecdotal. Moll and Legler (1971) reported that sliders in Panama were inactive at night, and slept at the edges of vegetational mats with the head and dorsal half of the carapace out of the water. Gibbons et al. (1990) noted that sliders seen at night in Georgia were normally underwater on the surface of the mud or buried beneath the substrate. Granda and Maxwell (1978) reported that captive sliders slept on the bottom of the tank, and about once an hour they opened their eyes, raised their heads to the surface, breathed, and then submerged again. Here I report observations on the sleep-like behavior of Bulletin of the Maryland Herpetologica! Society Page 39 Volume 29 Number 2 June 1993 young, captive red-eared sliders (T. s. elegans), and compare the behavior of T. scripta with that of G. elephantopus. Methods In March and April 1986, five young T. s. elegans (33-36 mm straight carapace length) obtained from a dealer were maintained indoors in a 38- liter aquarium during a 12: 12 light: dark photoregime, with the onset of light at 0700. The aquarium contained a rock providing terrestrial habitat for approximately 25% of the surface area; water 7-8 cm deep covered the remaining surface area. Water temperatures ranged from 18.3°C at night to 23.3°C during the day, and air temperatures ranged from 19.4-25.0°C. The turtles were fed freeze-dried Tubifexsp. nearly every morning. Three turtles fed regularly, and appeared to be in excellent health; the other two refused to feed and eventually died. Observations of sleep-like behavior were made randomly. In the absence of electroencephalographic data, I considered the turtles to be “asleep” when immobile with the eyes closed for 30+ sec, and “awake” when the eyes were open (cf. Flanigan, 1974; Flanigan et al., 1974; Hayes et al., 1992). During each observation I recorded for each individual turtle the substrate occupied (land or water) , whether or not the head and/or tail were extended, the number of legs extended (1-4), the vertical angle of the longitudinal body axis (< or > 45°), and the sleep state (“asleep” or “awake”) of the turtles. Observations were made during both light phases (light and dark); during the dark phase, observations were facilitated with the use of a red light. A total of 207 observations, 69 for each turtle, were made for each healthy turtle; data on the unhealthy turtles, which were virtually always “asleep,” were rejected. A minimum of 60 min elapsed between successive observations, and no more than two observations were taken within a 6 hr period; independence between observations was assumed. To determine if the three individual turtles differed in their behavior, I computed a series of three-way likelihood ratio chi-square tests by using log-linear models (XL2 statistic; Everitt, 1977). Each test examined the relationships between the individual turtles, their sleep state, and one of the other variables associated with sleep-like behavior (light phase, substrate choice, head extension, legs extended, tail extension, and body angle). The significance of the first-order interaction term (calculated as the difference in df and XJ between the main effects model and the independent addition of the first-order interaction term; see Everitt, 1977) indicated whether there was individual variation in behavior. When there was no significant individual variation in behavior, I assumed independence and combined data for all individuals. I then examined the relationships between variables using Page 40 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 conventional two-sample chi-square tests (X2 statistic; Siegel 1956). All statistical tests were computed with Statistix software (Heisey and Nimis, 1985); all probabilities are two-tailed, with alpha = 0.05. Results The significance of the first-order interaction terms between individual turtles and the variables associated with sleep-like behavior indicate that there was little individual variation; only with respect to tail extension did the turtles differ significantly in their behavior (Table 1). The consistent behavior of the turtles is illustrated by the observation that during the dark phase all three turtles were “asleep" 88.5% of the time (N=26 observations), while during the light phase all three turtles were “awake” 88.4% of the time (N=43). When the data for the three turtles were combined, they were “awake” 94.5% of the time during the light phase (N-129 observations), but only 5.1% of the time during the dark phase (N=78; X2= 159.54, df=l, P<0.00 1 ; see Table 2). During the light phase, the turtles were usually in the water (79.1% of the time, N=129); when in the water they were seldom “asleep” (4.9% of the time, N=102; see Table 2). When on land, the turtles were usually basking while remaining vigilant (i. e. , alert with the eyes open) , and were “asleep” only 7.4% of the time (N=27; Table 2). During the dark phase, the turtles were almost always in the water (96.2% of the time, N=78); when in the water they were almost always “asleep” (94.7% of the time, N=75; Table 2). During the dark phase, the turtles were nearly always “asleep” in the water (95.9% of the time, N=74), but were occasionally “asleep” on land (Table 2), When “asleep,” the head, legs and tail of the sliders were retracted more often than not, but the expression of these postures was highly variable (Table 3). The turtles often “slept” pressed up against the sides of the rock in a relatively vertical position, but also “slept” on the floor of the aquarium in a relatively horizontal position (with roughly equal frequency; Table 3). When “asleep” in water, the turtles always remained completely submerged, but when in the vertical position, the head was frequently close to the surface. The turtles assumed a more vertical position more frequently when “asleep” than when “awake” (Table 3). Discussion When considering the relative homogeneity of an aquarium environment and the small number or turtles in this study, it is not surprising that the individual turtles exhibited few differences in sleep-like Bulletin of the Maryland Herpetologicai Society Page 41 Volume 29 Number 2 June 1993 behavior. In a natural environment, greater individual variation in sleep-like behavior may be expected since microhabitat characteristics vary greatly. Aquatic turtles are adapted to aquatic environments and are capable of remaining submerged for long periods of time. Hence it is not too surprising that the turtles in this study preferred to sleep while submerged. The vigilant behavior exhibited by young P. scripta while basking contrasts with that of G. elephantopus, which usually rests in sleep-like repose when basking (Hayes et al., 1992). These differences may be due to differences in their antipredator strategies. When approached by a potential predator, G. elephantopus and other terrestrial tortoises generally withdraw their limbs and remain stationary, relying primarily upon their domed carapaces with small external openings for defense (Hayes at al., 1988; Hayes, 1989). While basking, P. scripta and other aquatic turtles usually flee into the water and hide when approached by a potential predator (e.g., Moll and Legler, 1971; Hayes, 1989). Because P. pseudemys flees into another medium when threatened by predation, remaining awake and alert while basking would seemingly be more advantageous than it would for G. elephantopus. However, G. elephantopus has few, if any, natural predators (Hayes et al., 1988). Adult P. scripta tend to flee more quickly than immatures, possibly because immatures rely more upon cryptic coloration and concealment to avoid predators (Moll and Legler, 1971). Both P. scripta and G. elephantopus usually basked with the head and limbs extended, and slept at night with the head and limbs retracted. Similar postures have been recorded in other species of terrestrial tortoises, and are usually associated with thermoregulatory behavior (e.g., Voigt, 1975; Douglas and Layne, 1978). Heat may be absorbed by aquatic turtles during periods of basking by extending the limbs and head, and may be conserved at night by decreasing the surface to volume ratio when the head and limbs are withdrawn, thus reducing heat loss by conduction and convection. These postural adjustments, apparently exhibited by both aquatic and terrestrial turtles, thus appear to have a common themoregulatoiy function. However, much more research is needed to better understand the ecological ramifications of behavioral sleep in turtles. Acknowledgments I thank K. R. Beaman and W. K. Hayes for assistance in finding literature. Page 42 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Literature Cited Ayala-Guerrero, F. 1987. Sleep in the tortoise Kinostemon sp. Experientia 43: 296- 298. 1988. Sleep patterns in a chelonian reptile (Gopherus Jlavomarginatus) . Physiol. Behav. 44: 333-337. Douglass, J. F., and J. N. Layne. 1978. Activity and thermoregulation of the gopher tortoise ( Gopherus polyphemus) in southern Florida. Herpetologica 34: 359-374. Everitt, B. S. 1977. The Analysis of Contingency Tables. John Wiley & Sons, Inc., New York. 128 pp. Flanigan, W. F. 1974. Sleep and wakefulness in chelonian reptiles: II. The red- rooted tortoise, Geochelone carbonaria. Archs. Ital. Biol. 112:253-277. _____ , C. P. Knight, K. M. Hartse, and A. Rechtschaffen. 1974. Sleep and wakefulness in chelonian reptiles: I. The box turtle, Terrapene Carolina. Archs. Ital. Biol. 112:227-252. Gibbons, J. W., J. L. Greene, and J. D. Congdon. 1990. Temporal and spatial movement patterns of sliders and other turtles. Pp. 201-215 in J. W. Gibbons (ed.). Life History and Ecology of the Slider Turtle. Smithsonian Institution Press, Washington, D.C. Granda, A. M., and J. H. Maxwell. 1978. The behavior of turtles in the sea, in freshwater, and on land. Pp. 237-280 in D. I. Mostofsky (ed.). The Behavior of Fish and Other Aquatic Animals. Academic Press, New York. Bulletin of the Maryland Herpetologica! Society Page 43 Volume 29 Number 2 June 1993 Hayes, F. E. 1989. Defensive stances in turtles. Herpetol. Rev. 20:4-5. ______ , K. R, Beaman, W. K. Hayes, and L. E. Harris, Jr. 1988. Defensive behavior in the Galapagos tortoise (Geochelone elephantopus )» with comments on the evolution of insular gigantism. Herpetologica 44:11-17. ______ , W. K. Hayes, K. R. Beaman, and L. E. Harris, Jr. 1992. Sleep-like behaviour in the Galapagos tortoise (Geochelone elephantopus). Herpetol. J. 2:51-53. Heisey, D,, and G. Niniis. 1985. Statistix: An Interactive Statistical Program for Microcomputers. NH Analytical Software, St. Paul, Minnesota. Pp unnumbered. Hermann, H., M. Jouvet, and M. Klein. 1964. Analyse polygraphique du sommeil de la tortue. C. R. Hebd. Seanc. Acad. Sci. (Paris) 258:2175-2178. Karmanova, I., and E. V. Churnosov. 1972. Electrophysiological study of natural sleep and wakefulness in turtles and hens. J. EvoL Biochem. Physiol. 8:47-53. Moll, E. O., and J. M, Legler. 1971. The life history of a Neotropical slider turtle, Pseudemys scripta (Schoepff), in Panama. Bull. Los Angeles Co. Mus. Nat. Hist. Sci. 11:1-102. Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-Hill Book Company, New York. 312 pp. Vasilescu, E. 1970. Sleep and wakefulness in the tortoise (Emys orbicularis). Rev. Roum. Biol. (Ser. Zool.) 15:177-179. Voight, W. G. 1975. Heating and cooling rates and their effects upon heart rate and subcutaneous temperatures in the desert tortoise, Gopherus agassizit Comp. Biochem. Physiol. 52A: 527- 531. Page 44 Bulletin of the Maryland Herpetologica! Society Volume 29 Number 2 June 1993 Table 1. Results of first-order interactions (from three-way log-linear analyses; see text) to determine whether variables associated with sleep-like behavior in Pseudemys scripta elegans differed between the individual turtles. Variables XL2 df P Individuals X Light Phase 0.00 2 1.00 Individuals X Sleep State 0.12 2 0.94 Individuals X Substrate Choice 3.83 f • 2 0.15 Individuals X Head Extension 3.02 2 0.22 Individuals X Legs Extended 14.02 8 0.08 Individuals X Tail Extension 8.24 2 0.02 Individuals X Body Angle 1.82 • \ 2 0.40 Table 2. Conditional cell totals for the sleep state and substrate choice of Pseudemys scripta during different light phases. The data for all turtles are combined. Light Phase Dark Phase Sleep State Land Water Land Water "Awake" 25 97 0 4 "Asleep" 2 5 3 71 Bulletin of the Maryland Herpetological Society Page 45 Volume 29 Number 2 June 1993 Table 3. Conditional cell totals for postural variables associated with sleep¬ like behavior in Pseudemys scripta, independent of light phase and substrate choice. The data for all turtles are combined for each postural variable except tail extension (see text). Variable "Awake" "Asleep" X2 df P Head Extended 116 31 66.7 1 <0.001 Retracted 10 50 Legs Extended Zero 5 31 One 9 13 TWo 6 18 83.96 4 <0.001 Three 7 5 Four 99 14 Tail of Turtle 1 Extended 40 20 4376 1 0.03 Retracted 2 7 Tail of Turtle 2 Extended 38 11 14.54 1 <0.001 Retracted 5 15 Tail of Turtle 3 Extended 37 10 20.34 1 <0.001 Retracted 4 18 Body Angle <45° 100 38 21.93 1 <0.001 >45° 26 43 V> Department of Natural Sciences, Section of Biology, Loma Linda University, Loma Linda, California 92350 Received: 9 December 1992 Accepted: 21 January 1993 Page 46 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 FIRST RECORD OF AMPHISBAENA DUBIA MULLER (REPT1LIA: AMPHISBAENIA) I N ARGENTINA A. Alberto Yanosky, James R. Dixon, and Claudia Mercolli One specimen of A mphisbaena dubia Muller was found at El Bagual Ecological Reserve, Northeastern Argentina, within a termite mound in a P asp alum grassland. This is the first record of the species in Argentina, previously only known from Brazil. Our locality is more than 900 km west of the Brazilian state of Santa Catarina, the southernmost limit for A. dubia. Both localities are at the same latitude (26°S). The specimen is clearly assignable to A. dubia and is unlike any of the nine species of Amphisbaena previously reported for Argentina. Se encontro un ejemplar de Amphisbaena dubia Muller en la Reserva Ecologica El Bagual, noreste de Argentina, dentro de uin termitero ubicado en un pastizal de P aspalum. Este es el primer registro de la especie en Argentina, previamente cononcida para Santa Catarina (Brasil), con su limite sur de distribucion a mas de 900 km de El Bagual. Ambas localidades se encuentran aproximadamente a la misma latitud (26°S). El especimen fue facilmente asignable a nivel especifico y difiere de las nueve especies de Amphisbaena previamente citadas para Argentina. Introduction One of our goals is an evaluation of the herpetofauna associated with terrestrial termite mounds within El Bagual Ecological Reserve (26° 10* 58” S, 58° 56’39" W). On 15 September 1991 an amphisbaenid was found while opening one of several termite mounds on the reserve . Attempts to determine the specimen’s identity made us conclude that it did not belong to any of the known species of Argentine amphisbaenids. The specimen was sent abroad for identification and all conclusions indicated that it was Amphisbaena dubia Muller. The genus Amphisbaena in South America has received little attention beyond Gan’s works of the 1960 and 1970’s. Particularly, A. dubia was described in 1924 by Muller from Piracicaba, Sao Paulo, Brazil, along with several new herpetozoa (Vanzolini 1978: 49). In 1964 Gans redescribed the species and added many new records that extended its range from the Brazilian state of Sao Paulo to the southern states of Parana and Santa Bulletin of the Maryland Herpetological Society Page 47 Volume 29 Number 2 June 1993 Catarina. The southernmost locality given by Gans (1964) was Tres Barras, Santa Catarina. The latter locality is approximately at the same latitude as El Bagual (26°S), but El Bagual is 900 km west of TYes Barras. Following Gans (1964), nothing else has been reported for this species. For Argentina, approximately nine Amphisbaena species have been cited (Peters and Orej as-Miranda, 1986). Amphisbaena darwinu A . camura, , A. fuliginosa, A. merteniu A . plumea, and A . prunicolor are characterized by a caudal autotomy, while our specimen does not have it. Amphisbaena steindachneri is characterized by more than 238 body annuli while our specimen has 23 1 , and finally A. alba and A. agustifrons , apart from being large and thick amphisbaenids, have more than 42 segments per midbody annulus, while our specimen has 35. Description El Bagual collection number RED-AD-001, female, total length 19.5 cm, snout-vent length 18.0 cm, body mass 2.0 g; collected by C. Mercolli & A. A. Yanosky, 15 September 1991. A medium sized amphisbaenid without fusions of head shields; two pairs of large parietals, right parietal divided with anterior larger, left parietal undivided; two rows of postgenials, first pair normal with no segment extending forward to contact the postmental; no postmalars; blunt tipped tail without autotomy constriction; no precloacal pores because of sex, but scale indentations suggest two pores per side; body annuli 215, from angulus oris to precloacals; 10 caudal annuli; color of preserved specimen light brown and pinky. The specimen is assigned to Amphisbaena dubia Muller because: 1 it has no major fusions of head shields, e.g. prefrontals, frontals, and supralabials in pairs; 2.- it is a medium sized specimen, usually with less than 35 (29-42) segments per midbody annulus; 3.- fewer than 232 body annuli; 4.- the tail is cylindrical without basal constriction; 5.- more than 185 body annuli; and 6.- tail tip is rounded, without a marked vertical keel (Gans, 1964; Peters and Orej as-Miranda, 1986). Our specimen differs from the redescription given by Gans ( 1964) , by having 1 9 caudal annuli instead of 1 7 ; no circular spot could be found in the center of each segment and the intercalated dorsal half- annulus was found in the four precloacal annuli instead of 10; many segments at midbody appear to be lightly divided and counts ranged from 32 to 35 segments with a mode of 35. There are three supralabials and two and one half infralabials. The second supralabial is the largest, with the first supralabial next in size. Page 48 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Coloration The specimen was slight violet and rosy when alive. Preserved, dorsal segments bear a light brown appearance. Each segment has a stronger brown coloration in the anterior part, and pigments disseminate at the end of each segment. All segments appear to be bordered by a creamish coloration which allows easy segment delineation. Head and segments of the first third of the body length appear more pigmented. Ventrally, segments bear pigmentation only on the anterior part, the remainder cream colored. The cream color extends the width of the belly over most of the ventrals, but is occasionally interrupted. Throat and chin cream colored. Habitat Gans (1964) did not present data on the habitat of this species. Our specimen was found in Paspahim grassland containing a scattering of palm trees (Copemicia alba), and within a termite mound that contained the ant genus Camponotus. The amphisbaenid appeared to be in a dormant state 30 cm deep in dissociated soils, not in the very abundant termite galleries. Amphisbaena darwini was found associated with vegetal residum in Acromyrmex ant mounds by Gallardo (1977). The grassland where our specimen was found is affected annually by two natural pulses, floods in summer-autumn, and fires in winter- spring, but termite mounds remain little altered after both occurrences. Distribution According to localities given by Gans (1964) Amphisbaena diibia appeared restricted to the Paranense Biogeographical Province of Cabrera and Willink (1973). This province occupies southeastern Brazil, Misiones in Argentina, and eastern Paraguay; an area supporting 1500-2000 mm of rainfall, with a marked dry season during winter and the terrain characterized by rolling hills. The species distribution now includes the humid chaco of northeastern Argentina with similar characteristics ( 1 200- 1 800 mm rainfall, marked dry season in winter- spring) , except the terrain is relatively flat. The species has not been found between 52-56°W longitude, but future surveys should confirm its presence there. Bulletin of the Maryland Herpetological Society Page 49 Volume 29 Number 2 June 1993 Literature Cited Cabrera, A.L. and A. Will ink. 1973. Biogeografia de America Latina. Serie Biologia OEA #13. Washington, D.C., 122 pp. Gallardo, J. M. 1977. Reptiles de los alrededores de Buenos Aires. EUDEBA ed., Buenos Aires, 213 pp. Cans, C. 1 964. Redescription of Amphisbaena dubia Muller (Amphisbaenia: Reptilia). BREVIORA 205:1-11. Peters, J. A. & B. Orejas-Miranda. 1 986. Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaeninas. Smithsonian Institution Press, Washington, D. C., 193 pp. Vanzolini, P. E. 1978. An annotated bibliography of the land and fresh water reptiles of South America ( 1 758- 1 975) . Vol.il. (1901-1975). Museu de Zoologia, Univ. Sao Paulo. 316 pp. El Bagual Ecological Reserve, Salta 994, 3600 - Formosa, Argentina (AAY; CM); Department of Wildlife & Fisheries Sciences, Texas A &M University, College Station, Texas 77843, USA (JRD). Received: 16 December 1992 Accepted: 21 January 1993 Page 50 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 A NEW SPECIES OF KNOBSCALE LIZARD (REPTILIA: XENOSAURIDAE) FROM MEXICO Hobart M. Smith and John B. Iverson The three species of Xenosauras ( X . grandts, X. newmanorum, X. platyceps) currently recognized (King and Thompson, 1968) occur in the tropical, mesic foothill regions on Atlantic slopes of Mexico and Guatemala, and on Pacific slopes in Oaxaca, Mexico. The discovery of another population on the Mexican plateau at as great an altitude as 2361 m (8400 ft), in a semiarid habitat, is of great interest. The first specimen collected of that population was a juvenile (now KU 140046) taken 8 May 1967, by Randy McCranie near Chapulco, southeastern Puebla. A subadult (now UF 41956) was later (21 Oct. 1970] taken by Fred G. Thompson 7.5 km (4.7) E Chapulco. Simulated by the realization that the latter specimen belonged to no known species of the genus, JBI and companions secured a series of 10 (UF 51438-47) on3May 1981, about 8 km (4. 7 mi) E Chapulco, at an altitude of about 2134 m (7000 ft), under flat rocks lying on other flat rocks on southeast-facing slopes (Fig. 1). The latter locality is off of Hy 150, 2.4 km (1.5 mi) southwest of the Veracruz/ Puebla state line, where the species was at that time relatively abundant. Subsequent study has corroborated the initial surmise that the population represented by the cited specimens is indeed different from any recognized at present and as reviewed by King and Thompson (1968). Comparisons of that population (herein referred to informally as the Chapulco population) with others has required use of characters not compared by King and Thompson (1968). Materials and Methods Comparisons of the 12 Chapulco specimens were made with 54 specimens of other taxa, listed in the Appendix, representing all three known species of the genus, and three (X. g. agrenon , X. g. grandis , X. g. rackhamtj of the five subspecies of X. grandis. No examples of X. g. sarunartinensis and X. g. arboreus have been examined. All material of the genus in the Florida Museum of Natural History (UF) and the University of Colorado Museum (UCM), and all in the University of Kansas Museum of Natural History (KU) except for nine X. g. grandis duplicates from Cuautlapan, Veracruz, and five X. newmanorum from Xilitla, San Luis Potosi, has been studied. Bulletin of the Maryland Herpetological Society Page 51 Volume 29 Number 2 June 1993 Characters we have found useful in differentiating the Chapulco population from others are as follows. 1. Nape mark. The most distinctive feature of the Chapulco population is the strictly transverse direction and midnape restriction of the nuchal band (Fig. 2). In all other populations of the genus, the nuchal band is V-shaped (e.g.. Fig. 3, of X. newmanorum) and extends posteriorly at least to the level of the foreleg insertions, and in some to the level of the axilla. It is often (regularly in X. platyceps) more distinct than any of the four following dark crossbands on body, especially in mature individuals. Only in rare, totally unicolor individuals (e.g., X. g. grandis , KU 52488) is the neck band not as described. The nuchal collar of the Chapulco species is wholly or partly interrupted by a whitish vertebral streak, but a similar light interruption occurs in all other taxa, although weakly in most X. g . grandis. The nuchal band is solid black (except for the vertebral interruption) only in juveniles of all taxa and most adult X. g. grandis. Otherwise, the band tends toward fragmentation, less pronounced on its borders, with maturation, much as is the case for the bands on the trunk, although usually less pronounced. 2. Ground color of dorsum. The second most distinctive feature of the Chapulco species is the whitish dorsal ground color on body and tail. All other populations of the genus have the entire dorsum pigmented; in them a narrow tan interval separates the five broad dark bands and that ground color extends elsewhere to the head, onto the tail between its dark rings, and to the lateral fold. Only the tips of the tubercles in the light areas of the Chapulco population are clearly pigmented (Figs. 2, 4). 3. Dark spots on venter. None whatever occur in the Chapulco species; the whitish ventral coloration extends dorsad past the lateral fold, whereas in all other taxa the darker dorsal ground color extends ventrad past the lateral fold onto the sides of the abdomen even in those (X. newmanorum , X. platyceps ) in which the venter itself is immaculate. All specimens examined of X. grandis have a spotted venter, at least on its sides (even in the dorsally unicolor example), although King and Thompson ( 1968) indicate that some X. g. agrenon do not have ventral dark spots. Those exceptions may represent other taxa or have the pattern obscured by premoulting changes of the epidermis. 4. Postparietal dark spots. In the Chapulco population, a pair of distinct, squarish dark spots is regularly present bracketing a vertebral light streakjust behind the head (Fig. 2). In some specimens these spots extend forward onto the rear parietal region. In no named population of the genus do such spots occur. Page 52 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 5. Postrostral The Chapuico population is unique in having a peculiarly bulbous, relatively large postrostral that is wider than long (1.3 to 2 times as wide as long; Fig. 4). In other populations it is usually smaller, less bulbous, and longer than wide; in some individuals it is not even recognizable. In no specimen we have examined, except in the Chapuico population, is it both large and wider than long. In one population (as yet unnamed, here termed the Yautepec population), it is usually split longitudinally. 6. Rostral In all specimens of the genus the rostral and supralabial region are peculiarly protuberant, especially as seen in dorsal view. The rostral is accordingly fiat-topped, except inX platyceps, which has a median posterior process of the rostral that is unique to and regularly present in specimens we have examined of that species. 7. Zygomaiic-postGcular ridges , The canthus temporalis varies considerably in the genus, its prominence largely dependent upon the degree to which the temporal tubercles are separated from each other by granular scales. Where there are no or few7 granules, and the tubercles are in contact with each other, the canthus temporalis is not clearly evident. A longitudinal row of scales is always present on the canthus, however, and curves anteroventrally posterior to the orbit, joining the postocular ridge (on the posterior border of the orbit) below the orbit. In the Chapuico species, the two ridges are tightly in contact throughout most or all of their parallel lengths (Fig. 4), whereas inX. g . grandis , X. g. agrenoix X. new mcu lomm an d X. platyceps they are well separated throughout their parallel lengths by X- 2 rows of flat scales, except where they meet below the orbit. X. g. rackhami and the Yautepec populations are like the Chapuico species in this character. 8. Ventral scale rows. We have found no interpopulational differences in number of transverse rows on the belly (counted posteriorly from midventer at axillary level to the preanal group of scales) or in the preanal region, or in a combination of these counts. However, the maximum transverse count is 25-29 in the Chapuico population, whereas in all other populations the variation is 18-24 (except for the unnamed Yautepec population, with 23-27). 9. Lamellae on fourth toe. The Chapuico population has 19-23 lamellae under the 4th toe, whereas X. g . grandis , X g. rackhami X newmanomm and X platyceps have 25-33. Only X. g, agrenon, with 21-29 lamellae, and the Yautepec population, with 19-24, are not separable from the Chapuico species bv this criterion. 10. Size and proximity of tubercles. Although not stressed by King and Thompson (1968), presumably because of subjectivity of evaluation. Bulletin of the Maryland Herpetologlcal Society Page 53 Volume 29 Number 2 June 1993 tubercle size and association are clearly of considerable taxonomic significance because of their marked interpopulational variation. X. platyceps (Fig. 5) has the most prominent tubercles, densely crowded over most of the dorsal surfaces of the body and limbs, as well as in the temporal region, except for the middorsal trunk region; the paravertebral tubercles are smaller than the lateral ones and do not form a paired row as in most other populations, in which the lateral tubercles on the trunk are smaller than or barely equal in size to the paravertebral tubercles. The distance between the lateral tubercles is to a considerable degree population-constant and interpopulation variable. For example, they are separated from each other by less than their own diameter in X. platyceps , by 1-1.75 times their diameter in the Chapulco species, and by 2-4 times in X. g. grandis . The distance between the flattened, relatively large tubercles in the paravertebral rows, where they exist, is of importance. Similarly, the tubercles on the dorsal surfaces of the thigh and brachium are in contact or separated by various numbers of rows of granules in different taxa. Also, the rear and posterodorsal temporal tubercles vary intertaxonomically in degree of separation from each other by granules. 1 1 . Tail/s-v proportion All populations of Xenosaurus have a tail/ s-v proportion varying a little above or below equality, except for the veiy short-tailed Yautepec population, and the veiy long-tailed population near Vista Hermosa, Oaxaca. 12. Supraorbital semicircles . In all populations of the genus the supraorbital semicircles are separated, except for infrequent narrow contact of one or two scales, with the exception of X. newmanorum , in which they are regularly (fide King and Thompson, 1968, and in the one specimen we have seen) in contact. 13. Supraoculo- orbitals. Between the supraorbital semicircles and the enlarged supraoculars lie one or two rows, depending on taxon, of moderately large “supraoculo-orbitals,” and also in some taxa a row of veiy tiny scales is present in addition to or in lieu of the moderate sized scales. The Chapulco series has but one row of moderate sized supraoculo-orbitals. 14. Labiomentals . In the Chapulco species, the labiomental series of scales always extends forward to the first chinshield and infralabial. In other taxa the series often ends at the 2nd or even the 3rd chinshield. 15. Sexual dimorphism . Females attain a somewhat greater size (expressed in s-v length) than males, apparently in all taxa of the genus. We have, however, found no clear-cut external difference between the two sexes. The base of the tail is somewhat wider in most preserved males, but not always; perhaps in life that difference is a more reliable indicator. Both sexes Page 54 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 have a peculiar, more or less rectangular postanal “platform” (Fig. 6) covering about six to eight scale rows; the round, posterolateral comers of that platform have their scales distinctly enlarged, so that those comers protrude from the surface of the tail. No differences in the extent of development of that platform or its protuberant rear comers are apparent in comparison of males and females. The one difference that appears to be constant is that the small scales toward the middle of the most posterior row of the platform are distinctly wider than long in males, and squarish in females. This difference is perhaps correlated with an otherwise rather imperceptibly wider base of the tail in males than in females, assuming that the number of scales does not differ. The difference in scale shape, even though probably requiring at least a hand lens to detect, may be useful to captive breeders. However, the observed difference is limited to the X. grcindis species group, including all members of the genus except X. newrnanonim and X. platyceps. The pertinent scales are squarish in both sexes of those two species. It should be noted also that inj ected specimens (rather than ones slit for preservation), of both sexes of all members of the genus (although to a greater extent in females), almost always have the cloaca partially everted, obscuring the postanal platform. Presumably this propensity is correlated with parturition of large neonates. Description In view of the numerous peculiarities of the Chapulco population, as previously detailed, it is regarded as a previously unrecognized species that we here name Xenosaurus rectocollaris sp. nov. Holotype. UF 51438, adult male, 2.5 km (1.5 mi.) SW Veracruz/ Puebla state line, Hy 150 (8 km (4.9 mi) E Chapulco), Puebla, Mexico, about 2134 m (7000 ft), 3 May 1981, John B. Iverson et al. Pamtypes. Eleven, including UF 51439-47, topotypes, same collectors and date; UF41956, 7.5 km (4.7 mi) E Chapulco, Puebla, Fred G. Thompson, 21 Oct. 1970; and KU 140046, nr Chapulco, Puebla, Randy McCranie, 8 May 1967. Diagnosis - Definition. A member of the genus Xenosaurus with nuchal collar straight across middle of neck, not V-shaped; venter without dark markings; light areas between dorsal dark bands whitish, tips of tubercles brownish; lateral fold whitish; a pair of squarish dark marks posterior to occiput; a single, large, bulbous postrostral, wider than long; zygomatic and postorbital ridges in contact through most or all of their parallel length; one row of moderately large supra oculo-orbitals; labiomental Bulletin of the Maryland Herpetological Society Page 55 Volume 29 Number 2 June 1993 row extending forward to 1st chinshield and infralabial; maximum number of scales in a transverse row of scales on venter 25-29; lamellae on 4th toe 19-23; most temporal tubercles contacting each other, no more than a few posterior tubercles separated by one row of granules; tubercles in paravertebral rows elong^'*, fiat, slightly larger than lateral tubercles, separated anteropo^teriorly from each other by one or two row-s of granules; paravertebral in . separated from each other by a distance varying from 2- 3 times the length of their tubercles; lateral tubercles of abdomen separated from each other by their own diameter, little more or less; dorsal foreleg and hind leg tubercles mostly in contact with each other, at most one row of granules between them; caudal dark bars solid black above, longer than light tan interspaces, but weakly split and narrower than light interspaces below. Description of hoiotype. As in diagnosis; postparietal dark spots dimly continued narrowly forward on either side of the midiine to frontal region; collar with two central light areas on either side, almost completely split vertebrally, extending laterally to level of upper edge of tympanum; four broad dark bars on trunk posterior to collar, each of the three crossing the dorsum with numerous small, light central areas, anterior and posterior borders intact, lateral ends open (as in the collar), reaching to but not onto lateral fold; sacral dark band nearly solid black, light central spots much smaller than in other trunk bands; 10 dark bands on tail, all solid black above except for a few veiy small, central light dots on the basal band; 4th and 5th caudal bands with a few central light dots below. Postrostral large, contacting both nasals; rostral straight -edged posteriorly; enlarged supraoculars 4-5, separated from supraorbital semicircles by a row of scales as large as most of those lateral to enlarged supraoculars; postorbital and zygomatic ridges in contact behind eye except for one large, dorsally intercalated scale; one row of lorilabial scales below suboculars, strongly keeled; 10-11 supralabials; 10-11 infralabials; labiomental row extending forward to 1st chinshield and 1st infralabial. Head length (straight line, to rear edge of tympanum) 23.5 mm, width 19 mm, depth 10 mm; s-v 95 mm; tail 94.5 mm; hind leg 38 mm. Variation Most significant variation is discussed under Materials and Methods. There are only two males in addition to the hoiotype: no. 51441, 87 mm s-v (tail 82 mm), and no. 41956, 78 mm s-v (tail 70 mm). The tail /s-v ratio varies from 0. 89 to 1 .06. In every specimen the labiomental row extends anteriorly to the 1st chinshield and infralabial. The tail bands vary from 9 to 13. The scales in the single row between the enlarged supraoculars and supraorbital semicircles are relatively large in all. The transverse rows Page 56 Bulletin of the Maryland Herpetoiogica! Society Volume 29 Number 2 June 1993 on venter, axillary level to preanal area, vary from 35 to 38, the preanal rows from 3 to 5. The smallest and largest specimens, both females, measure 45 (UF 51447) and 104 (UF 51440) mm, respectively. Etymology . The specific name rectocollaris is derived from the Latin rectus, straight, and coRare, collar, and is used as an adjective modifying the generic name. It alludes to the unique shape of the collar in this species, being straight transverse rather than v-shaped. In-as-much as the nuchal mark of other taxa of the genus has often been referred to as a collar, it is expedient that the specific name should designate kind of collar. Remarks. Color slides of X. rectocollaris reveal that the iris in this species is yellowish -orange, not bright red as inX. g. grandis from Cuautlapan, Veracruz. Prof. Jose L. Camarillo R. advises us (pers. comm.) that the iris is yellow in X. platyceps. One of the members of the 1981 field trip conducted by JBI, C. R. Smith, reported 28 August 1981 (in litt.), while a student at the University of Texas, that "while holding the Xenosaums , one gave birth to a single huge baby. On dissecting one for tissues in Michigan, I found a fairly advanced ovum... these one-lizard litters may be of interest,” especially since Fritts (I960) and Alvarez (1982) indicated a litter of size of four to seven in Cuautlapan X. g. grandis , three in Chiapas “X. rackhamC However, dissection of the five mature female para types of X. rectocollaris (84-104 mm s-v; miniatures 45-68 mm s-v) reveals one with two large eggs (15-17 mm) in one oviduct (the other oviduct missing at the present time, presumably having been removed for study), the remainder all with ovarian eggs only, the largest ones numbering 3-4 in each ovary and measuring 1.5-2 mm in diameter. It is rather surprising that such variation in embryonic development should exist among animals all collected on one day, from small ovarian eggs to large oviductal eggs to full parturitional maturity. At least it is obvious that the litters of X. rectocollaris are not limited to single neonates; Alvarez (1982: 133) provided a possible explanation for such a misimpression by the comment that in “X. rackhamf (translated) “...the females give birth to three young, not all at the same time but over a period of four to six days.” Other members of the herpetofauna sharing the semiarid habitat of X. rectocollaris , and found at the same time by JBI’s party in 198 1 , constitute a peculiar mixture of typically more mesic, forest - dwelling taxa (e.g. Sceloporus mucronatus aureolas , Crotalus l intermedins), as are other species of Xenosaums, and typically more xerophilic species {Sceloporus megalepidums pictus (subspecies fide Basmann and Smith, 1974), S, s. spinosus). The strictly plateau species Toluca lineata various was also found at the bottom of the hill where the Xenosaums were taken. Bulletin of the Maryland Herpetological Society Page 57 Volume 29 Number 2 June 1993 Acknowledgments We are much indebted to JBI’s field companions (Teny Leitheuser, Ron Magill, Peter A. Meylan, Paul Moler, C . R. Smith) for aid in collecting material at the type locality of X. rectocollaris\ to David L. Auth, Florida Museum of Natural History, as well as John Simmons, University of Kansas Museum of Natural History, and Dr. Shi-Kuei Wu, University of Colorado Museum, for permission to study material under their care; and to Dr. William M. Lewis, Chairman of the EPO Biology Department, University of Colorado, and Dr. Shi-Kuei Wu, for provision of vital facilities for research. Fieldwork in Mexico was supported by NSF DEB 8005586. The material from the type locality was collected under a permit issued to Peter Meylan by tire Direccion General de la Fauna Silvestre, Subdireccion de Aprovechamientos Faunisticos. Literature Cited Alvarez del Toro, Miguel. 1982. Los reptiles de Chiapas. Tercera edicion, corregida y aumentada. Tuxtla Gutierrez, Chiapas, Mexico, Institute de Historia Natural. 247 pp. Dasmann, Marlene and Hobart M. Smith. 1974. A new sceioporine lizard from Oaxaca, Mexico. Gr. Basin Nat., 34(3): 231-237. Fritts, Thomas H. 1 966. Notes on the reproduction of Xenosauras grandis (Squamata: Xenosauridae). Copeia, 1966(3): 598. King, Wayne and Fred E. Thompson. 1968. A review of the American lizards of the genus Xenosauras Peters. Bull. Florida St. Mus., Biol. Sci., 12(2): 93-123. Page 58 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Appendix Specimens examined of Xenosaurus , other than X. rectocollaris , are as follows. X g. grandis. VERACRUZ: Cuautlapan (KU 105840-5; UCM 48453- 6). X, g , agrenon. OAXACA: Santa Rosa, nr Lachao, District of Juquila (UCM 52518, 52607-8); Rio Sal, nr Lachao, District of Juquila (topotypes; UCM 41148-56, 44475-8). X g. rackhami CHIAPAS: Cerro del Sumidero, 10.7 mi N Tuxtla Gutierrez (UCM 19028). X. newmanorum. SAN LUIS POTOSI: 4. 1 mi E Xilitla, 2650 ft (UF 25006). X platyceps. TAMAULIPAS: 12.4 mi SW Rio San Marcos (Cuidad Victoria), Hw 101 (UF 42025-32). X sp. nov. OAXACA: Cerro Ac alt epee, San Juan Acaltepec, District of Yautepec (UCM 44463-74). X. sp. nov.? OAXACA: Vista Hermosa, 1600 m, municipality of Bulletin of the Maryland Herpetological Society Page 59 Volume 29 Number 2 June 1993 Fig. 1. The type locality of X. rectocollaris. 8 km E Chapulco, Puebla, Mexico. Page 60 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Fig. 2. A female paratype (UF 51440), 104 mm s-v, of X. rectocollaris. Bulletin of the Maryland Herpetological Society Page 61 Volume 29 Number 2 June 1993 Fig. 3. Xenosaurus newmanorum, UF 25006, male, 96 mm s-v, showing the large, V-shaped, posteriorly situated nuchal mark. Page 62 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Fig. 4. The same animal as in Fig. 2, showing the tight juxtaposition of zygomatic and postorbital ridges, and the large, median postrostral. Bulletin of the Maryland Herpetological Society Page 63 Volume 29 Number 2 June 1993 Fig. 5. Xenosaurus platyceps , showing the large and profuse tubercles on body and limbs. UF 42048 and 42032, both males, 106 mm s-v. Page 64 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Fig. 6. Xenosaurus platyceps, showing the nearly immaculate venter (marked only on the sides), and the postanal platform (arrow). UF 42025, female, 87 mm s-v. Bulletin of the Maryland Herpetological Society Page 65 Volume 29 Number 2 June 1993 Comaltepec (KU 87437; UCM 49320, 52482, 52487-8). Addendum Received too late to be incorporated into the preceding text, but here regarded as a paratype, is UF 86763, from 3 km NE Lagunas San Bernardino, Puebla, Mexico (within 20 km of the type locality), 18°36’06M N, 97°17’25,rW, 2500 m, found in a limestone rock pile 26 Oct. 1992 by E. L. Reiser and F. G. Thompson, It is half grown (67 mm s-v, tail 61 mm), male (not dissected, judged from scale shape as described in the text), and conforms fully with the diagnostic character-states of the species, including color and pattern; there are 26 (max.) contiguous ventrals across the abdomen, 34 rows axilla to groin, and the 4th toe lamellae are 23-23. Department of EPO Biology , University of Colorado , Boulder, Colorado, 80309-0334 (HMS); and Department of Biology, Earlham College, Richmond, Indiana, 47374 (JBI). Received: 15 January 1993 Accepted: 21 January 1993 Page 66 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 OBSERVATION AND REVIEW OF THE NESTING AND EGG-LAYING OF CORYTOPHANES CRISTATUS (IGUANIDAE). Marco A. Laze ano Barren) and Eleuterio Gongora-Arones The casque-headed iguanid lizard Corytophan.es cristatus is found from the state of Veracruz, Mexico to Colombia (Peters & Orej as-Miranda 1986; Villa, et a l 1988) inhabiting prime tropical rain forest (Stuart, 1948 and 1958; Duellman, 1963; Lieberman, 1986; Bock, 1987; Lazcano-Barrero, et al 1992) at elevations below 1000 m.a.s.l. (Johnson 1989; Campbell & Vannini, 1989). Due to its apparent low population-densities, arboreal habits, cryptic behavior, coloration and morphology (Davis, 1953; Stuart, 1958; Andrews, 1979; Bock, 1987), the ecology and life history of this species, as well as the other two members of the genus Corytophanes , remains poorly known, as indicated by Lang ( 1 989) in a recent review of the basciliscine iguanians. The only published information concerning nesting and egg laying has been that of Ream (1965) and Bock (1987). As part of an herpetofaunal inventory of the Selva Lacandona, Municipio de Ocosingo, Chiapas, Mexico (Lazcano-Barrero, et al., 1992), on July 27, 1984, an adult female C. cristatus (TL = 373 mm, SVL =119 mm) was collected, by MALE and EGA, while covering her hole nest (depth 54 mm; width 43 mm) in a slope (inclination 50 to 60 degrees) on a path (width ca. 40 cm) in a tropical rain forest, described by Breedlove ( 1973), at 1250 hrs., near Laguna Jalisco (16°44’N, 91°1 1W; 400 m elev.), after a prolonged (14 hour) and heavy rain. The nest contained a clutch of 7 eggs (Figure 1). The number and average measurements of these eggs (Table 1) are above those previously reported by Ream ( 1 965; N = 6; weight = 2.01 g; Length = 23.0 mm; Width = 12.4 mm) and Bock (1987; N = 5; length = 20 mm). The specimen (Figure 2), unfortunately lost, was deposited in the Instituto Nacional de Investigaciones Sobre Recursos Bioticos, herpetological collection (INIRB-035), at present under custody of the Instituto de Historia Natural del Estado de Chiapas. When detected, the female became cataleptic. Only after capture did the lizard become active, repeatedly attempting to bite the hand that held it, a behavior previously documented by Davis (1953) and Bock (1987). Bulletin of the Maryland Herpetological Society Page 67 Voiume 29 Number 2 June 1993 The lizard had dirt on top of its casque, supporting Bock’s (1987) observations that this structure is actively employed by females in nest excavation, a behavior also described in Corytophanes hemandezi (Perez- Higareda, 1981) the only other oviparous species of the genus. After being seized, the female defecated; the peculiar scat (Figure 3) contained remains of an orthopteran and a lepidopteran larva. These prey items agree with the specialized diet (large arthropods] reported by Andrews (1979) for this species in Costa Rica and Panama. Available information on the reproduction of C. crisiatus is summarized in Table 2, and indicates that the species nests on compact soils devoid of leaf litter, within prime tropical rainforests, laying 5 to 7 eggs. Although more eggs could perhaps be laid as evidenced by a specimen collected by Duellman (1965), which contained eight fully developed ova. A positive correlation between clutch size and female length (SVL) is also expected, as has been documented for other iguanid lizards (Ford & Seigel, 1989). The species nests between June and August, during the summer peak of the rainy season. According to Ream (1964), the known incubation period, under captive conditions, is five months for a single hatchling (TL =75 mm; SVL = 35 mm). Although detection of nesting females in rainforest trails may constitute a bias, at present nesting away from trails can not be determined from published sightings. Nevertheless it is clear that the species nests on open ground. We hypothesize that among rainforest iguanids, nest excavation on areas free of leaf litter may substantially reduce detection of females by potential predators, by eliminating the sound of leaf litter removal and reducing the time and energy required during the nesting excavation process, a period when the arboreal, cryptic, iguanid females of C. cristatus are more vulnerable to predators. Moreover, use of rainforest trails (compacted soil, no leaf litter) by C. cristatus may also reduce detection of eggs by terrestrial, leaf litter, and fossorial predators. Acknowledgments We thank Charles Duncan and Antonio Munoz-Alonso for reviewing the manuscript and Aureliano Montoya for printing the photographs. Fieldwork was possible through a grant (PCECBNA-02 1 189) from the Consejo Nacional de Ciencia y Tecnologia (CONACYT). Page 68 Bulletin of the Maryland Herpetologicai Society Volume 29 Number 2 June 1993 Literature Cited Andrews, R. M. 1974. The Lizard Corytophanes cristatus: an extreme “sit- arid- wait” predator. Biotropica 11(2): 136-139. Breedlove, D. M. 1 973. The phytogeography and vegetation of Chiapas, Mexico. Pp. 149-165, in Vegetation and vegetational history of northern Latin America (A. Graham, ed.) Elseiver Sci. Publishing Company, Amsterdam. Bock, C. B. 1987. Corytophanes cristatus nesting. Herp. Review 18(2):35. Campbell, J. A. & J. P. Vamiini. 1989. Distribution of amphibians and reptiles in Guatemala and Belize. Proceed. West. Found. Vert. Zool. 4(1): 1-21. Davis, D. P. 1953. Behavior of the lizard Corytophanes cristatus. Fieldiana, Zool. 35:1-8. Duellman, W. E. 1963. Amphibians and Reptiles of the rainforest of Southern el Peten Guatemala. Univ. Kansas Publ. Mus. Nat. Hist., 15:205-249. Ford, N. B. and R. A. Seigel. 1989. Relationships among body size, clutch size, and egg size in three species of oviparous snakes. Herpetologica 45(1) :75- 83. Johnson, J. D. 1 989. A biogeographic analysis of the Herpetofauna of northwestern Nuclear Central America. Milwaukee Publ. Mus. Contrib. Biol. Geol., 76:1-66. Lang, M. 1989. Phylogenetic and biogeographic patterns of bascilicine iguanians (Reptilia: Squamata: “Iguanidae”. Bonner Zoologische Monographien 28:1-171. Bulletin of the Maryland Herpetologica! Society Page 69 Volume 29 Number 2 June 1993 Lazeana -Barrera, M. A., E. Gongora-Arones & R. C. Vogt, 1992. Anfibics y reptiles de la Selva Lacandona. in: Vasquez- Sanchez, M. A. y M. A. Ramos (eds.). Reserve de la Biosfera Montes Azides, Selva Lacandona: Investigacion para su Conservacion. Publ. Esp. Exosfera 1: 145-171. Lieberman, S. S. 1986. Ecology of the leaf litter herpetofauna of a neotropical rain forest: La Selva, Costa Rica. Acta Zool. Mex. (ns), 15: 1-72. Perez-Higareda, G. 1981. Nesting and incubation times in Corytophanes hemandezi (Lacertilia: Iguanidae). Bull. Maryland Herp. Soc. 17(2): 71- 73. Peters, J. A. & B. Orej as -Miranda. 1 986. Catalogue of the Nepotropical Squamata. Part II Lizards and Amphisbaenians. Smithsonian Institution Press, Washington D. C. 1-293. Ream, C. H. 1965. Notes on the behavior and egg laying of Corytophanes cristatus. Herpetologica 20: 239-242. Stuart, L. C. 1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. Publ. Mus. Zool. Univ. Michigan 69:1-109. 1958. A study of the Herpetofauna of the Uaxactun-Tikal area of northern El Peten, Guatemala. Contrib. Lab. Vert. Biol. Univ. Michigan 75:1-30. > . Villa, J., L. D. Wilson and J. D. Johnson. 1988. Middle American Herpetology a bibliographic checklist. University of Missouri Press. Columbia. 1-131. Page 70 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 TABLE 1 WEIGHTS AND MEASUREMENTS OF Corytophanes cristatus EGGS. EGG No. WEIGHT LENGTH WIDTH g mm mm 1 2.2 22.0 13.0 2 2.6 23.0 13.2 3 2.7 23.5 14.0 4 2.6 24.0 13.5 5 2.6 23.5 13.0 6 2.8 24.0 13.9 7 2.7 24.6 13.0 X 2.6 23.5 13.4 TABLE 2.- SUMMARY OF REPRODUCTIVE INFORMATION OF C. cristatus . FEMALE SVL (mm) CLUTCH SIZE DATE/ ACTIVITY HABITAT LOCALITY SOURCE 125 (a) JUNE 28 RAINFOREST CHINAJA ALTA VERAPAZ GUATEMALA DUELLMAN, 1965 120 5 (b) JUNE 25 NESTING TRAIL PRIMARY RAINFOREST RARA AVIS HEREDIA COSTA RICA BOCK, 1987 n. d. n. d. n. d. NESTING TRAIL PRIMARY RAINFOREST LA SELVA COSTA RICA BOCK, 1987 119 7 JULY 27 NESTING TRAIL PRIMARY RAINFOREST SELVA LACANDONA CHIAPAS MEXICO PRESENT 107 6 FEB 5(c) NESTING CAPTIVE 3ARRO COLORADO PANAMA REAM, 1964 a. - Eight ova within female, with an average diameter of 11.1 mm. b . - The clutch size may have been higher since the female was captured and released on site, without determining if there were additional eggs in the oviduct. c . - Eggs laid in captivity by a female that had been isolated for five months and 10 days, mating took place before August 27 the previous year when the specimen was captured. Bulletin of the Maryland Herpetological Society Page 71 Volume 29 Number 2 June 1993 Page 72 Bulletin of the Maryland Herpetological Society Eggs of C. cris talus, (photographed by MALE). Volume 29 Number 2 June 1993 Bulletin of the Maryland Herpetological Society Page 73 Volum© 29 Number 2 June 1993 Page 74 Bulletin of the Maryland Herpetological Society Volume 29 Number 2 June 1993 Program for Studies in Tropical Conservation & Department of Wildlife and Range Science, University of Florida, Gainesville Florida 32611 0304. Current address: ECOSFERA A. C./ Centro de Investigaciones Ecologicas del Sureste, Apartado Postal 219, San Cristobal de Las Casas, Chiapas 29200, Mexico (MALE); Departamento de Fomento Pesquero, Presidencia Municipal de Champoton, Calle 23 it 17, Champoton, Campeche 24400, Mexico (EGA). Received: 25 January 1993 Accepted: 20 February 1993 Bulletin of the Maryland Herpetological Society Page 75 Volume 29 Number 2 June 1993 ‘MOVES THE MID-ATLANTIC REPTILE SHOW Captive Born Reptiles (only), Accessories, & Books for sale PROCEEDS WILL PURCHASE RAINFOREST This Is A Non - Profit Venture By The MB Herp Society September 25th & 26th 1993 Saturday 10 a.m. to 5 p.m. Sunday 9 a.m. to 5 p.m. Door Prize Drawings! Marriott‘s Hunt Valley Inn off 1-83 Just North of Baltimore MD Hotel Reservations (Show Discount): Phone # 800-228-9290 or 410-785-7000 M-F 8-6 Admission Donation: Two Day Pass Adults $10.00, Children (under 12) and Seniors $7.00 (Additional Donations Welcome) Lectures Saturday evening 7-11 PM: Keynote Speaker-DR. ROGER CONANT Sandy Barnett, Jack Cover, Wayne Hill-Additional speakers to be announced INFORMATION AND VENDOR RESERVATIONS Contact : The Maryland Herp Society, 2643 N. Charles St. Balto. MD 21218 Phone: (410) 235-6116 Wednesdays 9-5 only or Show Coordinator Tim Hoen - Phone (410) 557-6879 anytime * No venomous reptiles ‘ Not responsible for damage, theft, or injuries * No mammals or insects Page 76 Bulletin of the Maryland Herpetological Society Society Publication Back issues of the Bulletin of the Maiyland Herpetological Society, where available, may be obtained by writing the Executive Editor. A list of available issues will be sent upon request. Individual numbers in stock are $2.00 each, unless otherwise noted. The Society also publishes a Newsletter on a somewhat irregular basis. These are distributed to the membership free of charge. Also published are Maryland Herpetofauna Leaflets and these are available at $.2 5 /page. Information for Authors All correspondence should be addressed to the Executive Editor. Manuscripts being submitted for publication should be typewritten (double spaced) on good quality 8 1/2 by 11 inch paper with adequate margins. Submit original and first carbon, retaining the second carbon. If entered on a word processor, also submit diskette and note word processor and operating system used. Indicate where illustrations or photographs are to appear in text. Cite all literature used at end in alphabetical order by author. Major papers are those over 5 pages (double spaced, elite type) and must include an abstract. The authors name should be centered under the title, and the address is to follow the Literature Cited. Minor papers are those papers with fewer than 5 pages. Author’s name is to be placed at end of paper (see recent issue). For additional information see Style Manual for Biological Journals (1964), American Institute of Biological Sciences, 3900 Wisconsin Avenue, N.W., Washington, D.C. 20016. Reprints are available at $.03 a page and should be ordered when manuscripts are submitted or when proofs are returned. Minimum order is 100 reprints. Either edited manuscript or proof will be returned to author for approval or correction. The author will be responsible for all corrections to proof, and must return proof preferably within 7 days. The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc. 2643 North Charles Street Baltimore, Maryland 21218 Maryland Herpetological Society US ISSN: 0025-4231 AT^ 93bulletin of the TFtarylanb f)erpetoIogical ©ociety DEPARTMENT OF HERPETOLOGY THE NATURAL HISTORY SOCIETY OF MARYLAND, INC. MdHS . A FOUNDER MEMBER OF THE Eastern Seaboard Herpetological League 30 SEPTEMBER 1993 VOLUME 29 NUMBER 3 BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY Volume 29 Number 3 September 1993 CONTENTS Dromicus giganteus J an (Reptilia: Serpentes) is a Nomen Nudum Hobart M. Smith, James R. Dixon and V. Wallach . . 77 The Date of Publication of Jaltris Cope (Reptilia: Serpentes) Hobart M. Smith and V. Wallach . . . 80 A Review of the Members of the Sceloporus variabilis Lizard Complex Hobart M. Smith, Gonzalo Perez-Higareda and David Chiszar . 85 The Generic Allocation of Tantilla canula (Reptilia: Serpentes) Hobart M. Smith, Oscar Flores Villela and David Chiszar . . . . . 126 The Status of the Members of the Sceloporus aeneus Complex (Reptilia: Sauria) of Mexico Hobart M. Smith, Jose L. Camarillo R. and David Chiszar . 130 Notes on Hibernation of the Smooth Green Snake Opheodrys ve malts, in New Mexico James N. Stuart and Charles W. Painter . 140 The Maryland Herpetological Society Department of Herpetology Natural History Society of Maryland, Inc. 2643 North Charles Street Baltimore, Maryland 21218 BULLETIN OF THE Volume 29 Number 3 September 1993 The Maryland HerpetotogicaS Society Department of Herpetology, Natural History Society of Maryland, Inc. Executive Editor . Herbert S. Harris, Jr. Steering Committee Frank Groves Jerry D. Hardy, Jr. Herbert S. Harris, Jr. Library of Congress Catalog Card Number: 76-93458 Membership Rates Membership in the Maryland Herpetological Society is $16.00 per year. Foreign $20. 00/year. Make all checks payable to the Natural History Society of Maryland, Inc. Meetings The third Wednesday of each month, 8: 15 p.m. at the Natural History Society of Maryland (except May — August, third Saturday of each month, 8:00 a.m.). The Department of Herpetology meets informally on all other Wednesday evenings at the NHSM at 8:00 p.m. C' tK ' ' ' ; Volume 29 Number 3 September 1993 DROMICUS GIGANTEUS JAN (REPTILIA: SERPENTES) IS A NOMEN NUDUM Hobart M. Smith, James R. Dixon and V. Wallach Although treated by Dixon (1981, 1989) as an available name having priority over Dromicus omatus Garman (1887) (now Liophis omatus ), Dromicus giganteus Jan (1863) is a nomen nudum and therefore has no nomenclatural status. No action by the International commission on Zoological Nomenclature is necessary to protect the name Liophis omatus (Garman). In commendable deference to the long-established goal, in response to a vital need, of the International Code of Zoological Nomenclature to safeguard nomenclatural stability-a deference that should be more widely pract ic e d- D ixon (1981) chose to maintain existing nomenclatural usage when, in monographing the eastern Caribbean snakes of the genus Liophis, he discovered that a never-used name he considered available, Dromicus giganteus Jan (1863:67), antedated the long- accepted name Dromicus omatus Garman (1887:281). Under those circumstances, the security of the latter name, in the combination Liophis omatus since Dixon’s 1981 review, would rest strictly with the willingness of other workers to resist application of the Principle of Priority that, under the Code, would require use of Jan’s name unless the International Commission on Zoological Nomenclature was requested to intercede through its plenary powers. On the contrary, any nomenclatural fundamentalist, herpetologist or not, could insist upon using the earlier name. So far that has not happened. For example, the most recent monograph of the West Indian herpetofauna (Schwartz and Henderson, 1991 :625) still uses Carman’s name. However, such principled maintenance of the status quo is by no means assured, hence a reexamination of the situation was undertaken. Much to our relief, a critical evaluation of Jan’s 1863 proposal of Dromicus giganteus , in the light of the 1985 edition of the Code, leaves no question that it actually is a nomen nudum , and therefore is not available in nomenclature. Article 12(a) explicitly requires that to be available any new scientific name published before 1931 “must have been accompanied by a Bulletin of the Maryland Herpetological Society Page 77 Volume 29 Number 3 September 1993 description or a definition of the taxon that it denotes m (italics ours). True enough, Jan (1863:66-67) gave a partial key to the members of Dromicus, but D. giganteus was in a group of nine species, all having a single anterior temporal, 17 scale rows and one loreal. The species of that group were not distinguished from each other, hence there is no characterization of that particular taxon (D. giganteus)-ohfy to a group of species for which no collective discriminatory name was given. Since D. giganteus is clearly a nomen nudum and unavailable under the Code, the specimen on which it was based is not a holotype. Article 12(c) makes it clear that mention of a specimen does not comply with the requirement of Article 12(a). Furthermore, under Article 11(e) Dixon’s interpretation of D. giganteus as available, and his characterization of it, does not make that name available even of his authorship because he did not adopt it as a valid name. Literature Cited Dixon, James R. 1981. The Neotropical colubrid snake genus Liophis: the eastern Caribbean complex. Copeia, 1981:296-304. 1989. A key and checklist to the Neotropical snake genus Liophis with country lists and maps. Smithsonian Herp. Inf. Serv., (79): 1-40. Garman, Samuel. 1887. On West Indian reptiles in the Museum of Comparative Zoology at Cambridge, Mass. Proc. Am. Philos. Soc., 24:278- 286. Jan, Georges. 1863. Elenco sistematico degli ofidi descritti e disegnati per l’lconografia generale. Milano, Italy. 143 pp. Schwartz, Albert and Robert W. Henderson. 1991. Amphibians and reptiles of the West Indies. Gainesville, Florida, xvi, 720 pp. Page 78 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Department of Environmental, Population and Organismic Biology, University of Color ado, Boulder, Colorado 80309-0334 (HMS); Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station , Texas 77843 (JRD); and Department of Herpetology, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 (VW). Received: 1 February 1993 Accepted: 20 February 1993 Bulletin of the Maryland Herpetological Society Page 79 Volume 29 Number 3 September 1993 THE DATE OF PUBLICATION OF JALTRIS COPE (REPTILIA: SERPENTES) Hobart M. Smith and V. Wallach The earliest publication date of the nominal genus Jaltris , an unjustified emendation of Ialtris Cope, 1862, is in Cope (1863), not in any of several other works as has been averred in various publications. As an available name, although currently invalid, Jaltris Cope has precedence over any other generic name of identical spelling and later date, and could be used as a replacement name if needed. The generic name Jaltris has long been of uncertain attribution and date, although universally accepted as an emendation, valid or not, of Ialtris Cope (1862: 73). Although Cope used the spelling Jaltris in several publications following the appearance of the volume containing his description of Ialtris (1879, 1886, 1887, 1894, 1895, 1 900) , he never cited the source of the emendation, leading Cochran (1941) to attribute Jaltris to his earliest publication using that name (Cope, 1879). In 1900 Cope actually used both spellings: Ialtris on p. 698, presumably a lapsus, and Jaltris on p. 1091, but otherwise he consistently used the latter spelling in reference to its biotaxon. Marschall (1873: 50) has commonly been accepted as the first explicitly to list Jaltris as an emendation of Ialtris Cope, but again without giving a source. Therefore Scudder (1882b: 163) assumed that Marschall created the emendation, although in error; Scudder erred also in placing Jaltris in the Lepidoptera, although he listed Ialtris Cope (1862) correctly in both works (1882a: 171; 1882b: 159). His action led Schulze et al. (1932: 1707) and Williams and Wallach (1989: 79) likewise to credit Marschall (1873) with the emendation. Neave (1939-1950), oddly, does not list Jaltris. Outside of literature-recorders, Werner (1924: 131) is the only author except Cope to have used Jaltris as a valid name, but again without citation of source. Although never so noted before now. Cope himself created the emendation. It is found in the “Errata and Addenda” section of volume 14 of the Proceedings of the Academy of Natural Sciences of Philadelphia - the same one in which his description of Ialtris appeared. In that section, p. 594, is the entry “p. 73, line 34, for Ialtris read Jaltris .” Page 80 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Original spellings changed in publisher’s “corrigenda” (Art. 33(b)(1)) are, under Art. 32(c)(ii), incorrect original spellings, and as such are to be corrected (Art. 32(d)), the replaced name then having no separate availability as of the corrected work. That interpretation would, in the present case, make Jaltris the correct name for the genus universally (since 1924) known as laltris , and the latter name unavailable as of Cope (1862). On the contrary, however, Art. 32(c)(ii) requires that, for that rule to apply, corrigenda must appear “in the original publication itself,” which Jaltris Cope (1863) did not do. The section of the Proceedings containing Cope’s original description of laltris, according to Nolan (1913: xiii), appeared at some time before May 27, 1862, whereas the section containing the “Errata and Addenda” appeared at some time between Jan. 12 and Feb. 26, 1863. Accordingly, under the current Code, Jaltris actually is an unjustified subsequent emendation of laltris, originating with Cope (1863). As such it is an available although currently invalid name. It has precedence over any other generic name of identical spelling and later date, and could be used as a replacement for laltris were it needed, e.g. through discovery of junior homonymy. Had the two names been published in the same work it would have been a different story. However unfortunate this outcome may be from the standpoint of Cope’s intent, it is most fortunate relative to nomenclatural stability, inasmuch as the spelling laltris has been regularly accepted, with the cited exceptions, at least since Garman (1887: 284), in dozens of works by dozens of authors. We here note another citation of Jaltris that is of ambiguous significance. As pointed out by Williams and Wallach (1989: 79), Troschel (1863: 635), in his listing of the herpetological literature of 1862, cited “ Jaltris Cope nov. gen. Proc. Philadelphia p. 72,” taken by Williams and Wallach as an “Error or emendation of laltris Cope.” Since only works published in 1862 were covered in this compilation, and only p. 72 was cited, it is quite possible that Troschel himself independently created the emendation. Even if so, it clearly must have appeared long after Cope’s emendation, as indicated by the early publication of the latter (between Jan. 12 andFeb. 26, 1863), and the probable late appearance of Troschel’s article (suggested by its advanced pagination, pp. 620-641). On the other hand, because of the late date of appearance of the latter, Troschel may indeed merely have adopted Cope’s emendation without citing its source precisely (p. 594). The issue of which alternative is correct is moot, however, since Cope’s emendation can be accepted as having appeared earlier than Troschel’s work. Bulletin of the Maryland Herpetological Society Page 81 Volume 29 Number 3 September 1993 Acknowledgment We are much indebted to Dr. Philip K. Tubbs, Secretary of the International Commission on Zoological Nomenclature, for critical counsel and authorization of the interpretation here propounded. Literature Cited Cochran, Doris M. 1941. The herpetology of Hispaniola. Bull. U. S. Natl. Mus., (177): i-vii, 1-398. Cope, Edward D. 1862. Synopsis of Holcosus and Ameiva , with diagnoses of new West Indian and South American Colubridae. Proc. Acad. Nat. Sci. Philadelphia, 14: 60-82. 1863. IJaltris.] In Errata and addenda. Proc. Acad. Nat. Sci. Philadelphia, 14: 594. 1879. Eleventh contribution to the herpetology of tropical America. Proc. Acad. Nat. Sci. Philadelphia, 18: 261-277. 1 886. An analytical table of the genera of snakes. Proc. Acad. Nat. Sci. Philadelphia, 23: 479-499. 1887. Catalogue of batrachians and reptiles of Central America and Mexico. Bull. U. S. Natl. Mus., (32): 1-98. 1894. The classification of snakes. Am. Nat., 28: 831-844. 1895. The classification of the Ophidia. Trans. Am. Philos. Soc., 18: 186-219. 1900. The crocodilians, lizards and snakes of North America. Rep. U. S. Natl. Mus., 1898: 153-1270. Garman, Samuel. 1887. On West Indian reptiles in the Museum of Comparative Zoology, at Cambridge, Mass. Proc. Am. Philos. Soc., 24: 278-286. Page 82 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Marschall, Augusto de. 1873. Nomenclatorzoologicus .... Vindobonae, Carol! Uebexreuter, iv, 482 pp. Neave, Sheffield Airey. 1939-1950. Nomenclator zoologicus. 5 vols, London, Zoo!, Soc. London. Nolan, Edward J. 1913. An index to the scientific contents of the Journal and Proceedings of the Academy of Natural Sciences of Philadelphia, 1812-1912. Philadelphia, Acad. Nat. Sci. Philadelphia, xiv, 1419 pp. Schulze, F. E., W. Kuekenthal, and K. Heider. 1932. Nomenclator animalium generum et subgenerum. Vol. 3. Berlin, Preuss. Akad. Wiss. Pp .1299-2248. Scudder, Samuel H. 1882a. Nomenclator zoologicus. I. Supplemental list of genera in zoology. Bull. U. S. Natl. Mus., (19): i-xix, 1-376. 1882b. Nomenclator zoologicus. II. Universal index to genera in zoology. Bull. U. S. Natl. Mus., (19): i-xix, 1-340. Troschel, F. H. 1863. Bericht ueber die Leistungen in der Herpetologie waehrend des Jahres 1862. Arch. Naturg., 29: 620-641. Werner, Franz. 1924. Uebersicht der Gattungen and Alien der Schlangen der Familie Colubridae. II. Teil (Dipsadomorphinea und Hydrophiinae). Arch. Naturg., 90A: 108-166. Williams, Kenneth L., and Van Wailach. 1989. Snakes of the world. Vol. I. Synopsis of snake generic names. Malabar, Florida, Krieger, viii, 234 pp. Bulletin of the Maryland Herpetological Society Page 83 Volume 29 Number 3 September 1993 Department of EPO Biology, University of Colorado, Boulder, Colorado 80309-0334 (HMS); and Department of Herpetology, Museum of Comparative Zoology , Harvard University, Cambridge, Massachusetts 02138 (VW), Received: 26 February 1993 Accepted: 10 March 1993 Page 84 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 A REVIEW OF THE MEMBERS OF THE SCELOPORUS VARIABILIS LIZARD COMPLEX Hobart M. Smith, Gonzalo Perez- Higareda and David Chiszar In the absence of convincing evidence of intergradation with Sceloporus uariabilis , and because of near parapatry and sympatry, S. teapensis is regarded as a full species. It is found to be extensively sympatric with S. smithi, which is therefore also regarded as a full species. S. v. olloporus is revived for the subspecies extending from Guatemala to Costa Rica. The range of S. teapensis splits that of S. v. variabUis into three completely dichopatric enclaves : (1) southern Tamaulipas to southern Veracruz; (2) the coast of Tabasco; and (3) Pacific slopes of Oaxaca east of the Isthmus ofTehuantepec through southern Chiapas and southwestern Guatemala. S. smithi evidently hybridizes occasionally with S. teapensis, and may well do so also with S. v. variabilis in southeastern Oaxaca. Several characters not previously explored are surveyed, and a key to the five recognized taxa of the complex is provided. Urophely (a new term) is reported for the first time in the Sceloporinae. The correct taxonomic rank for the taxon first named Sceloporus teapensis by Gunther in 1890 has been uncertain throughout most of its history. Boulenger ( 1 897) accepted it as a valid species, although erroneously regarding Bocourt’s (1874) S. cupreus as an earlier name for it. Even then Boulenger noted that it might be referable to S. variabilis Wiegmann (1834), presumably as a distinct subspecies. Its more recent history begins with maintenance as a full species, under its proper name S. teapensis, by Smith (1936, 1939), but suggestions later by various workers that subspecific rank under S. variabilis might be correct culminated with a formal adoption at that level by Cole (1978), based on morphological and karyological studies of the entire variabilis group, although the misinterpretation of a critical series from Tabasco was a major factor in his conclusion. In 1982 Sites and Dixon supported Cole’s conclusion in a highly sophisticated analysis of external variables in 1408 specimens representing the entire species S. variabilis as they then understood it. The same arrangement appeared in Mather and Site’s (1985) review of that species in the American Catalog, and most subsequent works have conformed with that arrangement, in which four subspecies are recognized; S. v. marmoratus Hallowell (1852), S. v, smithi Hartweg and Oliver (1937), S. v. teapensis and S. v, variabilis . Smith’s (1937, 1939) analyses differed not only in ranking S. teapensis as a full species but also in segregation of the southern-most enclave of S. variabilis , Bulletin of the Maryland Herpetological Society Page 85 Volume 29 Number 3 September 1993 in Central America, as a separate subspecies, S. v. ollopoms Smith (1937). Although the review by Sites and Dixon (1982) involved a total of 38 characters, their taxonomic conclusions were based largely upon two essential characters: dorsal scale count and number of femoral pores. They did not consider number of postrostrals, and found no basis for special emphasis on presence of absence of a subnasal (utilized by Smith (1939) in differentiating S. teapensis from S. variabilis ) or number of longitudinal rows of dorsals. Guillette et al. ( 1 983) reported an additional character of possible use in distinguishing the latter two taxa - pigmentation of the testicular peritoneum. The present study was undertaken to assess the utility of these variables to the understanding of the relationships of the taxonomic populations of the S. variabilis complex, with special emphasis upon “S. teapensis Gunther”. Materials and Methods A total of 1384 specimens was examined, without screening for sample size, and in part selected with special attention to any evidence of geographic range overlap of taxonomically distinct populations. In that context even single specimens from given localities can be of great significance. All specimens of the complex were examined in the collection of the University of Kansas Museum of Natural History (KU; 686) and the University of Colorado Museum (UCM: 355); and selected specimens in the American Museum of Natural Histoiy (AMNH; 5), University of Michigan Museum of Zoology (UMMZ; 6), Museo de Zoologia, Facultad de Ciencias, U.N.A.M. (MZFC; 25), Field Museum of Natural History (FMNH; 74), University Of Illinois Museum of Natural (UIMNH; 54), Museum of Comparative Zoology (MCZ; 2), James Ford Bell Museum of Natural Histoiy, University of Minnesota at Minneapolis (JFBM; 216), and the University of Texas at Arlington Vertebrates Collection (UTA; 151) (acronyms from Leviton et al., 1980). In addition. Dr. Jack Sites kindly made available to us all of his data on 1408 specimens, many not duplicated in our materials, especially the 486 in the Texas Cooperative Wildlife Collection (TCWC). The KU, UCM, and UTA collections constituted the heart of our study; other specimens were for the most part selected to clarity specific questions in critical areas. Inasmuch as Sites and Dixon (1982) dealt thoroughly with the geographic variation in dorsal scale counts and number of femoral pores (their Figs. 2, 3), we concluded that nothing could be gained by repetition of that work, although both character-states were recorded on certain critical specimens. In addition, we agreed that data on most other characters are “of little utility in assessing patterns of geographic variation” (Sites and Dixon, Page 86 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 1982: 15). In several characters, however, some in their list and some not, we found significant geographic variation, as follows. 1 . Nuchal (nape) rows. The sizes of the dorsal, ventral and lateral scales on the body in Scelopoms are, to a considerable degree, proportional. Certainly their sizes are not as independent of each other as in Anolis and its allies, but neither are they exactly correlated. The dorsal body scale size, however, appears to vary as a unit, and thus in the S. variabiiis complex is reflected more or less equally well by ( 1) the usual head- tail count and by (2) a transverse count at those points on the body where dorsals and laterals are sharply differentiated from each other. Two such points exist: in the nuchal and sacral regions. In both regions, the most lateral dorsal scales are distinctly larger than the adjacent laterals, and are not basically uniplanar but describe an obtuse angle marking the junction of dorsal and lateral body surfaces. The nuchal (dr nape) count, as we recorded it, is the minimum transverse count between head and foreleg level, usually near the middle of the neck, and including the most lateral row of angular dorsals on each side. The range of variation in the complex is from 9 (S. teapensis ) to 18 (S. v. marmoratus ), forming a taxonomically stepped S-N cline, few rows to numerous ones, between those two taxa, but a N-S cline elsewhere. 2. Sacral (rump) rows. Similarly, as we recorded it, the sacral (or rump) count is the minimal count made transversely of the enlarged dorsals at the level of the hind legs. The dorsal scale rows converge in the sacral region, hence it is necessary to limit the area where the minimal count is made. The transverse line counted could not, in our procedure, include any scale lying in any part at or posterior to the midline between the anterior and posterior levels of the hind leg insertion. Total variation in the complex is 7 (S. teapensis ) to 15 (S. marmoratus). 3. Lateral abdominal scales. The scales that we have observed to vary in size inter-populationally, more or less independent of the size of the dorsal scales, are those on the sides of abdomen and neck. Unfortunately we have found no way to quantify the differences observed, hence they remain subjective and useful primarily in cases in which objective criteria leave doubt of identity. In S. teapensis the scales anterior to the immediate groin area are relatively large , little reduced in size compared with the median lateral scales (Fig. 1). In S. variabiiis and S. smithi, on the contrary, those scales are very small, little larger than those at the groin (Fig. 2). Bulletin of the Maryland Herpetological Society Page 87 Volume 29 Number 3 September 1993 4. Lateral nuchal scales. The posteroventral free edge of the lateral nuchal fold bears enlarged, acuminate, keeled scales contrasting sharply in size and shape with the scales lining the lateral nuchal pocket which is covered by the lateral nuchal fold. In S« teapensis the external scales between the series on the edge of the lateral nuchal fold and the dorsal nuchal scales are of about the same size as those on the edge of the fold, not reduced in size (Fig. 3) . They are little smaller than the scales in the lateral row of dorsal nuchals. On the contrary, in S. variabilis and s. smiihi the lateral nuchals are markedly smaller than the dorsal nuchals or the scales on the edge of the lateral nuchal fold (Fig. 4). 5. Lateral fold. The median lateral scales in S. smithi are so small- little larger than the scales in the borders of the axillary and groin regions- that, at least in preserved specimens, the larger scales dorsal and ventral to the small median ones tend to encroach upon the latter, raising the skin bearing the small scales into a longitudinal ridge (Fig. 5). All preserved specimens checked for this character possess the ridge at least weakly developed. Whether it is evident in live specimens is unknown. In other related taxa no such ridge occurs; the median lateral scales are presumably too large to form a ridge, and there is a greater contrast in size between them and the border axillary and groin scales. The lateral ridge appears to be structural, because it is positioned and shaped the same in all specimens. It lies rather high on the sides, distinctly nearer the dorsum than the venter, does not reach the groin, but curves ventrad anteriorly to end in the axilla close to the ventral insertion of the foreleg. It disappears completely with tension on the skin ventral to it, but its invariable position and shape suggests that it depends on some histological irregularity presumably of genetic origin. It does not occur even in S. v. marmoratus , which has equally small body scales. 6. Canthosubnasals. In the S, variabilis complex, either two or three scales extend along the canthal ridge, from the superciliaries to below the nasal scale. In previous works (e.g. Sites and Dixon, 1982), it was assumed that if only two scales are present along the entire ridge, it is the subnasal that is absent, because often it is extremely small where three scales are present. However, the subnasal in some specimens is large, even when three occupy the ridge. It is not certain that reduction of the total to two is always a result of loss of the subnasal rather than of the anterior canthal. We therefore here adopted the policy of referring to all of the scales collectively as canthosubnasals, of which there always were two or three. S. Page 88 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 teapensis typically has only two, varying from 61 to 86% in different populations (x 74%), whereas no other populations of the S. varkzbilis complex has two in more than 14% except for S. v . marmoratus with 31%. Care has to be taken to exclude from the canthosubnasal count any of the lorilabials (loreolabials of Stuart, 1971), which form a continuous series from below the subocular to the nasal. 7. Postrostrals. As pointed out long ago by Smith (1937), the number of postrostral scales in populations of Nicaragua and Costa Rica is often reduced to two from the normal four. We explored variation in that number throughout the complex, because it is a character often constant in a whole group; two regularly occur, for example, in members of the scaiaris and siniferus group, whereas four is typical in most other groups of the genus. Some care has to be taken to avoid confusion of the postrostrals with the anterior lorilabials. The total variation found in this complex is 2-6; counts of 5 or 6 are rare and anomalous. 8. Dorsal pattern . The dorsal pattern is essentially the same throughout the complex, with the exception of S. smithL In all populations except the latter, the ground color is gray-brown, with a pair of dorsolateral stripes, not sharply delimited, vaiying from a little to distinctly lighter than but of much the same hue as the ground color. A series of dark spots or marks parallels each light stripe near its medial edge, and on the sides of the abdomen near the lateral edge of the stripe. Juveniles and females exhibit more distinct spotting than adult males. In S. smithL on the contrary, the area between the dorsolateral stripes is nearly or quite uniform, dark chocolate brown or black; the dark spots of other populations are not or but faintly evident. The two light stripes are sharply defined and brilliantly distinguishable, essentially white in color, not dark-pigmented. The pattern is one of striking contrast individually and in comparison with other populations of the complex; it occurs in both sexes and at all ontogenetic stages. 9. Semeions . The gular and abdominal semeions (color patches; Smith et al, 1991) vary to such an extent that we have observed no populational differences, except for occurrence of weak abdominal semeions in adult female S. smithL They are rarely evident in S. v. variabilis. Bulletin of the Maryland Herpetological Society Page 89 Volume 29 Number 3 September 1993 10. Tail color. The small juveniles of S. smithi have a conspicuously light, pink tail, which at maturity attains a coloration like that of the body. Presumably the pink tail serves much the same function - diverting the attention of predators from a vital to a non-vital part - as in many s kinks. It is, however, the only instance of such urophely (Greek oura, tail, and phelos, deceptive) known to occur in Sceloporus or any other Sceloporinae. 1 1 . Carinal ridges. In S. smithi the keels on the dorsal trunk scales are particularly well defined and so nearly perfectly aligned that they form conspicuously continuous ridges. They are not equally as prominent in other populations of the complex. 12. Size. S. v. marmoratus is by far the smallest taxon of the complex, reaching a maximum s-v length of only 58 mm, and infrequently reaching 50 mm or more (16%) . All other taxa reach a maximum somewhat exceeding 70 mm; 78 mm is the maximum record (S. smithi) . \ 13. Testicular peritoneum pigmentation. Guillette et al. (1983) recorded that no testicular pigmentation was found in 50 male S. teapensis , whereas it was observed to occur asymmetrically only on the left testis in 139 (35%) of 398 male S. variabilis representing all subspecies. However, the absence of pigmentation in 65% of S. variabilis, and its disappearance outside of the breeding season, render the character virtually useless taxonomically. It was not considered important in our study, and was not recorded. Results Overview. As indicated in the accompanying key, all of the characters precedingly listed, except for no. 13, exhibit distinct geographic variation importantly supplementing the already well- established geographic variation in dorsal scale count and number of femoral pores. Utilizing all of these features, few problems were encountered allocating individuals or local populations to their proper taxon as here recognized. Initially, our primary concern was the taxonomic rank of Gunther’s S. teapensis, because of its histoiy of treatment as both a separate species and a subspecies of S. variabilis. All other taxa described in the past as subspecies of S. variabilis were initially assumed to be correctly accepted or Page 90 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 rejected at the level as concluded by Sites and Dixon (1982). As the study expanded little by little, it was therefore a great surprise to discover that at least one more of the subspecies, other than S. teapensis (S. smith!) , is definitely a separate species, and that S. v. olloporus is a valid subspecies. Nevertheless, as indicated by the accompanying map (Fig. 5), S. teapensis by its central position geographically holds the key to the entire complex. Its range splits the populations commonly referred (e.g. Sites and Dixon, 1982) to S. v. variabilis into three parts, here termed enclaves, apparently completely separated from each other. The Northern Gulf enclave extends from southern Tamaulipas approximately to the Rio Papaloapan in lowlands (farther south in uplands, at least to the Sierra Juarez). In the lowlands southeastward S. teapensis replaces S. v. variabilis , and extends to the coast eastward about to the Rio Coatzacoalcos, thence across the base of the Yucatan peninsula to Belize, and southward to central Guatemala on the east, northeastern Oaxaca on the west, including southern Tabasco and northern Chiapas. The southern central and southwestern parts of the Isthmus of Tehuantepec is occupied by S. smithl The ranges of those two species completely separate the Northern Gulf enclave of S. v. variabilis from the populations here called the COG enclave (an acronym for Chiapas, Oaxaca and Guatemala) , which is limited to Pacific slopes of Oaxaca east of the Isthmus of Tehuantepec, southern central and northeastern Chiapas, and southwestern Guatemala. The third isolated population, here termed the Southern Gulf enclave, was first discovered, so far as we are aware, by Dr. Thomas H. Fritts in 1967 (although first reported by Cole, 1978). Itisbyfarthedistributionally smallest of all taxa and enclaves of the S. variabilis complex, being limited, so far as is now known, to coastal Tabasco, north of the inland range of S. teapensis. The specimens Cole (1978: 8) reported of this enclave were referred by him to S. v. teapensis, on distributional grounds; we are confident, however, that they represent S. v. variabilis, since they agree in dorsal scale count (49-51) with his counts for that subspecies, and not with his counts (36-47) for S. teapensis . Playa Paraiso, where Cole’s specimens came from, is bracketed by the large series we have examined from Tabasco, all clearly S. v. variabilis. Ramirez and Gonzalez (1991) reported reproductive data on this enclave, properly identified. Although it seems highly unlikely, these three enclaves may actually have continuity, inasmuch as at least limited sympatry with S. teapensis is now known to occur. The subspecific taxon S. v. olloporus that we here revive ranges from eastern Guatemala south of Peten to Costa Rica, intergrading with S. v . variabilis in southwestern Guatemala and adjacent Chiapas. Bulletin of the Maryland Herpetologica! Society Page 91 Volume 29 Number 3 September 1993 Observations on each of these seven taxa and enclaves follow. 1 . Scelopoms smitht This taxon is arguably the most highly differentiated and distinctive of the S. variabilis complex, at least compared with sympatric and adjacent members. It is immediately recognizable at all postembryonic stages in both sexes by its distinctive pattern, having a very dark, chocolate brown to black ground color on back and sides, without or with very dim dark spots, and brilliant white, wide (2 and 2 half- rows at widest point) dorsolateral stripes. No other taxon of this complex has such a pattern, and no other is distinguishable by pattern. The dorsal scale count is higher (54-69, X 62-63, fide Hartweg and Oliver, 1937, and Sites and Dixon, 1982) than in any other taxon in the complex except S. v. marmoratus (54-72, fide Sites and Dixon, 1982: 19). The small size of the body scales is reflected in both taxa by the large number of rump (11-14, X12, and 11-15, X 12.5, respectively) and nape (13-17, X 15, and 14-18, X 16, respectively) rows, with means higher than in any other populational segments here treated of the complex. The rump rows in both taxa are 12 or more in at least 85% of the specimens examined, but in no more than 22% (range 0-22) of all other populational segments. The highland populations of Hidalgo and Queretaro, as noted by both Smith (1939) and Sites and Dixon (1982), referred to S. v. variabilis , also have small body scales, similar to those of S. v. marmoratus and S. smith , but we have not dealt here with their taxonomic status, having seen but five specimens of them. They may well constitute a distinct race. Other peculiarities of S. smithi include the extraordinary pink tail-tip of juveniles, previously mentioned; no other Sceloporus exhibits such urophely. The striking alignment of the keels of the dorsal scales, forming conspicuously continuous ridges, is also unique. The species holds the present record of maximum s-v length (78 mm, UCM 39859) in the complex, although both S. v. variabilis and S. teapensis may reach the same size, with a recorded maximum of 74 mm for the latter (Boulenger, 1897) and 76.4 mm for S. v. variabilis (UIMNH 10927, Tonala, Chiapas, fide data kindly provided by Dr. Jack Sites; two other specimens in the same series, UIMNH 10938- 9, measured 75.4 and 75.9 mm s-v). S. v. olloporus reaches much the same maximum, at 74.8 mm (KU 85916). Even the peculiar Queretaro population of S. v. variabilis approaches the maximum, at 73.9 mm (TCWC 29763). Hartweg and Oliver ( 1 937) noted that females of S. smithi are of much the same color as males “except that the colors are in general less intense”, and that their abdominal semeions are “not so distinctly bordered, and they exhibit veiy little of the pinkish hue.” Nevertheless, females of S. smithi regularly exhibit distinct abdominal semeions, whereas they are not evident at all in female S. teapensis, and seldom in s. variabilis. Page 92 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 A quite subjective difference between S. smithi and its adjacent taxa (S. teapensis and S. v. variabilis) is the reduced size of the central lateral abdominal scales, and the concomitant existence of a longitudinal ridge extending from near the lower margin of the axilla along the upper side of the abdomen to near the groin (but not into it). So far as we are aware, that ridge is unique to the species. S. teapensis and S. smithi represent the extremes of body scale size in the entire complex and also of pattern. Therefore it is not surprising that they are extensively sympatric. Reproductive isolation is no doubt at least reinforced if not largely dependent upon visual interspecific discrimination, given the sharp distinction in coloration between the two species. Sympatiy of these two species in the same localities is documented by several UCM specimens, all collected by that legendary naturalist Thomas MacDougall, who, although primarily a horticulturist, collected many vertebrates for the American Museum of Natural History, the University of Illinois Museum of Natural History, the University of Colorado Museum and the University of Kansas Museum of Natural History. Three S. teapensis (39849-51) and one (39852) S. smithi were taken near Palomares, Oaxaca, a few kilometers northeast of an area well represented in various museums by the former species (Tollosa, Tollocito, Real de Sarabia, La Princesa, etc.). As indicated on the accompanying map, however, S. smithi has long been known in nearby localities, as in the Juchitan region. There is no reason to doubt its occurrence there. MacDougall also obtained three of the latter species (38957-9) and one S. teapensis (39848) at Tamasola, district of Tequisixtlan, Oaxaca, a locality northwest of Tehuantepec and almost in the middle of the range of S. smithi All sympatric specimens are completely typical of their taxa. Although clearly no intergradation exists between S. teapensis and S. smithi two specimens (KU 43733-4), here assigned to the latter species, appear to be hybrids with S. teapensis. Both are adult females and have the scutellation typical of S. smithi (dorsals 68-70, rump rows 14-14, nape rows 17-16, canthosubnasals 3-3, lateral folds present), but their dorsal pattern is typical of S. teapensis. They were taken 2 mi N, 6 mi W Nejapa, Oaxaca, well within the range of S. smithi (see Fig. 6). We can interpret these specimens only as hybrids; they are the only evidence of which we are aware that interbreeding ever occurs between S. teapensis and S. smithi It is not equally apparent the S. smithi does not intergrade with, and is therefore not a subspecies of, S. v. variabilis , of which a population (COG) is isolated in southern Chiapas and immediately adjoining areas to the west in Oaxaca and to the east in Guatemala. Typical 8. smithi occurs as far Bulletin of the Maryland Herpetological Society Page 93 Volume 29 Number 3 September 1993 southeast as Ixtepec, Oaxaca (FMNH 1472(7)), and typical S. v. variabilis (COG) as far west as Niltepec, Oaxaca (FMNH 1468, 99251-2, UIMNH 21954), localities only 53 air km apart. It remains uncertain, however, how these two taxa interact between these two localities, if indeed their ranges actually meet, as seems likely. Hartweg and Oliver (1937: 4) claimed that “There is a specimen from Tapanatepec, Oaxaca (UMMZ 78852), which appears to be an intergrade between the two subspecies; it retains the coloration of smithi but exhibits the scale formulae of variabilis Smith (1939: 281), referring apparently to the same specimen, stated that it “tends strongly toward v. variabilis , having a dorsal scale count of 55. The dorsal coloration, however is identical with that of v. smithi We have concluded, however, that some mixup of data has been responsible for this erroneous record of an intergrade. Smith did not list any specimen from Tapanatepec under “S. v. smithi ', and the only ones from there under S. v. variabilis were two MCZ specimens (33454-5). We have re-examined those two, as well as four others from nearby (KU 37791, 2 mi E Tapanatepec; UTA 5674, Cerro Baul, 22 km N Tapanatepec; UIMNH 37356, 37264, Plan de Campana, Rio Porto Moneda, 30 km NNEN Tapanatepec, on Gulf slopes), and all are typical COG S. v. variabilis in both scutellation and coloration (although KU 37791 does have a distinct lateral fold). The so-called “intergrade” cannot be found. Greg Schneider kindly verified Jan. 30, 1992, that UMMZ 78852 was cataloged with eight specimens; seven are now present, all S. siniferus obtained in exchange with MCZ; the identity and whereabouts of the missing, eighth specimen is unknown. Jose Rosado kindly reported Febr. 5, 1992 (in letter) that indeed seven specimens are missing from the original series of 19 duplicates of S. siniferus entered under MCZ 28326-75. However, one specimen from the tagged series is missing, although its loose tag remains in the jar. It now appears impossible to verify the record of an intergrade from Tapanatepec, and we conclude that it should be disregarded until confirmed with either new material from that area or rediscovery of the specimen originally reported. On the other hand, it is quite likely that the specimen referred to is actually UMMZ 88404 from Arriaga, Chiapas, only some 32 km ESEE of Tapanatepec, since it is cataloged as S. v. smithi Greg Schneider kindly recorded the following data on it: a poorly preserved juvenile, 25 mm s-v, with 52 dorsals, dorsal coloration very dark, dark spots scarcely visible, dorsolateral light lines well defined and as much as three scales wide. More material from that area would be required for definitive conclusions. The locality is so distant from verifiable records of S. smithi that we suspect discoloration is responsible for its resemblance to that species. We accordingly here regard it as representing S. v. variabilis, although it may possibly (although we think it unlikely) represent a hybrid between the two taxa. Page 94 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 In support of the intergradation concept. Smith (1939: 281) also stated that “Specimens of v. variahilis from El Hule, Oaxaca, tend somewhat toward v. smithi in coloration”. However, El Hule is nowhere near the range of S. smithi, it lies on Atlantic slopes just south of the Veracruz border and just north of 18° N latitude (see Fig. 6), and the specimens from there cannot be influenced by the S. smithi genome. There simply is insufficient material available at the present time to demonstrate conclusively whether intergradation occurs between S, smithi and S. v. variahilis . Apparently that material should be sought in the area between Ixtepec and Niltepec, Oaxaca. Certainly since S. smithi and S. teapensis can hybridize, interbreeding between the former and S. v. variahilis should be possible. Nevertheless the very distinctive coloration of S. smithi should be highly effective in interspecific discrimination by the lizards themselves, maintaining their separate identities. Furthermore, S. teapensis and S. variahilis appear to maintain their separate identities, in spite of a greater superficial similarity than exists between S. smithi and S. variahilis , where their ranges are closely approximated and even interdfgitated in Guatemala and on Atlantic slopes of southern Veracruz and Tabasco (see following discussion). We accordingly conclude that S. smithi is allospecific to both S. teapensis and S. variahilis:; that relationship to the former is well assured, but to the latter requires confirmation. 2 . Sceloporus teapensis. The taxonomic validity of S. teapensis , at least as a subspecies of S. variahilis , as never been refuted. Its primary claim to validity has always been its large scales on body, 36 to 49, whereas its adjacent relative, S. v. variahilis, usually has 50-61. Although most data in Sites and Dixon ( 1 982) for S. teapensis conform approximately with those of Smith (1939) and Cole (1978), with an upper limit of 50 (the counts by Sites and Dixon extended to the level of the anus instead of the rear margin of the thighs, hence typically included 1-2 scales more than counts by the other workers), they cite a maximum of 58 in their series from the vicinity of La Libertad, El Peten, Guatemala. That series, however, included at least three S. chrysostictus (KU 55805, 55810, 55825) with 47, 50 and 53 dorsals (their counts). One other specimen (KU 59714), in the same series, they reported with 54 dorsals, a figure that cannot be verified because that specimen has now been skinned, cleared and stained; since other specimens of S. chrysostictus were accepted as S. variahilis and had comparably high counts, however (ranging in the species 42-57, fide Smith, 1939), and all other confirmed S. teapensis in their data, from that locality, had 42-50 dorsals, we conclude that KU 59714 also is a S. chrysostictus. One other specimen, UMMZ 74972, reported with 58 dorsals, actually has 43 or 44, Bulletin of the Maryland Herpetological Society Page 95 Volume 29 Number 3 September 1993 hence 58 is a clerical error. Thus 49 remains (by the thigh-level count) as the usual maximum for S. teapensis. Nevertheless, considerable overlap of even this restricted range of dorsal scale counts of S. teapensis occurs with S. v. variabilis , according to the data in Sites and Dixon (1982), and also our own data. The lowest count for S. v. variabilis in Sites and Dixon (1982) is 43, occurring in a series from Playa Miramar (1 1 km SW Front era), Tabasco. We have examined the same series, and the lowest count obtained in it was 45 (UIMNH 87391). Since the methodology of Sites and Dixon should yield higher, not lower counts, we assume that the count of 43 is a lapsus for some higher number. Even with a minimum dorsal count of 45 in S. v. variabilis , the overlap with S. teapensis is considerable, 45-49. More important, the nearest samples of S. v. variabilis to the range of S. teapensis , as given by Sites and Dixon, to the west in Veracruz (19 km N Alvarado and 8 km N Lerdo de Tejada), and to the east in Tabasco (Sanchez Magallanes and Playa Miramar) have means more closely approaching those of S. teapensis than any others of the subspecies west of Chiapas and Guatemala. Our own data on number of rump and nape rows, as well as these rows totaled (see Table 1) reveal strong but incomplete distinction between S. teapensis and S. v. variabilis , the former usually having lower counts than the latter. The samples from Tabasco and Alvarado areas of S. v. variabilis more closely approach S. teapensis in these respects than do others of their subspecies, paralleling the trends in dorsal scale counts. The reduced number of canthosubnasals (2-2) is also characteristic of S. teapensis , as compared with S. v. variabilis with 3-3, although to a somewhat lesser degree. However, again the frequency of occurrence of two is lowest in samples of S. teapensis from Oaxaca and Veracruz, which lie near the range of the northern populations of S. v. variabilis , as compared with the samples elsewhere, even though in Guatemala the two taxa occur in some areas in near parapatry or in sympatry. Although S. teapensis usually has fewer rump and nape rows, and fewer canthosubnasals, comparisons of the sums of all three with the same figures for S. v. variabilis gives no better separation of the two taxa than the dorsal scale count (see Table 1). An apparent documentation of sympatry of the two taxa exists in the form of one S. variabilis (UTA 20752) and seven S. teapensis (UTA 20333- 5, 20748-51) cataloged from Finca La Perla, ElTesoro, El Quiche, Guatemala, Jan. 9, 1986, J. A. Campbell. Campbell has confirmed (pers. comm.), however, that the single S. variabil is was not collected by him there, although the others were; the S. variabilis was given to him by a friend who had Page 96 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 recently come from the southern highlands of Guatemala, where that species is well known; its exact origin is uncertain, but is definitely not the same as the seven S. teapensis . It presumably represents S„ v. oUoporus , having 20 femoral pores. Although no documentation now exists of syxnpatry of the two taxa in Guatemala, they are recorded in close proximity in several areas in various works by L. C, Stuart, who never reported either any difficulty in distinguishing them, or populations that appeared intermediate. Most impressive is his 1943 report of 10 S. teapensis from Fihca San Francisco (“forty-one miles east and slightly north of Huehuetenango”), virtually surrounded by records for S. variabRis. We likewise have found no intermediates in Guatemala or Chiapas, and none is evident in those areas in Sites and Dixon's (1982) revision. Most recently, Campbell and Vannini (1989) summarized the ranges of the two species in Guatemala by faunal areas; their conclusions support ours. They note occurrence of only S. teapensis in the Peten area, and only 8. vartahilis in the Zacapan, Jalapan and Fuegan areas; both are recognized in the Quecchian area and the Cuchumatan subarea of the Huehuetenangan area. Range overlap and interdigitation are likely in both of the latter two regions. It is thus our conclusion that to the south, where S. teapensis extends into Oaxaca on the west, and in Chiapas and Guatemala elsewhere, it is sympatric or parapatric with S. smithi (Oaxaca) and 8. variahUis (both southern subspecies) with no evidence of intergradation.. Whether the species occurs in extreme northwestern Honduras is unknown, but it is unlikely since J onathan Campbell has not found it in the Puerto Barrios area (pers. comm.). Indeed, no representatives of the S. variahilis complex has been found between Lago de Izabal/Rio Dulce and the Honduras border. If present, they presumably would represent 8. v. oUoporus. On the contrary, the contact zones with the Northern Gulf and Southern Gulf enclaves of 8. v. variabRis give clear indication of the mutual influence of one taxon on the other. Introgression has apparently occurred in both directions, but whether it continues at present is uncertain. The rather narrow zones of intermediacy suggest that the contacts are secondaiy and that interbreeding is infrequent. However much the means approach each other toward the lines of parapatry, no sample yet reported is really ambiguous of allocation.The nearest geographic approach yet sampled of S. v. variabilis to S. teapensis is Sites and Dixon's (1982) series from 9 km NE of Lerdo de Tejada, Veracruz, and on the basis of dorsal scale count most of that series (± 75%) falls with 8. v . variabRis. We have not seen that series, but Bulletin of the Maryland Herpetological Society Page 97 Volume 29 Number 3 September 1993 another (JFBM 2852, 2858, 2860-1, 2865) from veiy close nearby (2 mi S Tlacotalpan, 1; 1.8 mi S Buena Vista, 4) is definitely referable to S. v. variabilis (all with 3-3 canthosubnasals, rump rows 10-11, nape rows 13- 14). No coastal or near-coastal samples are known to us from the presumed actual contact zones of S. teapensis and S. v. variabilis; a hiatus of some 60 km or more, whence specimens of neither taxon are known, exists to both the east and the west. Whether intergradation or hybridization occurs in those zones is uncertain; variational overlap elsewhere suggests that but one intermediate population occurs, but sympatry of two separate populations is not ruled out. The most critical series of specimens we have examined in this context is the one from Veracruz designated WNGS in Table 1. Those 39 include the ones already mentioned from Buena Vista and Tlacotalpan, and 35 as follows: coast road 20 mi SE Veracruz [= 48 km NW Salinas], BCM 45615; 15 mi NW Alvarado [= 2 km SE Salinas], UCM 50767; beach 5 mi SE Salinas [= 17 km NW Alvarado], UCM 39460-72; 4 km SE Alvarado, KU 59706-10; and Alvarado, KU 26736-7, 26740, 26742-3, 26752, 26754, 26756, 26761, 26764-6, 27037, 44383. All of these 35 represent S. v. variabilis , but there is a strong tendency toward the characters that define S. teapensis , viz. rump rows 8 in one, 9 in six, 10 in twenty-one, and 1 1 in seven; nape rows 11 in four, 12 in two, 13 in fourteen, 14 in ten, 15 in three, 16 in one. The canthosubnasals are 2-2 in two, 2-3 in one, 3-3 in thirty-six. As indicated in Table 1, the number of rump rows in the 8-9 range, characteristic of S. teapensis, is distinctly more frequent of occurrence in the southern part of the Northern Gulf enclave of S. v. variabilis (nearest the range of S. teapensis) than elsewhere in that species (18% vs. 0-7%). Hie same is true of the nape rows; 1 1 or fewer (common in S. teapensis) occur in 1 1% of the WNGS group, as opposed to 0-3% in other groups of the species. The canthosubnasals are, however, more or less typical of S. v. variabilis. No reciprocal tendency to have more numerous rump or nape rows is evident in the group of S. teapensis (Table 1, TOV) nearest the range of the Northern Gulf enclave, but the proportion with three (as in most S. v, variabilis) instead of two (as in S. teapensis) canthosubnasals is distinctly higher (39%) than elsewhere in the species (14%). Thus the variation in the three most useful characters (rump and nape rows, certainly reflecting the size of the dorsal scales and the dorsal scale count, and the canthosubnasals) distinguishing these two taxa in southern Veracruz suggests rather strongly that they are not sympatric, as they may well be in Guatemala, but that they hybridize or intergrade there, or did so until relatively recently. Page 98 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 However, what really happens where the two taxa meet in southern Veracruz is unknown because there is a gap of 50-60 km or more between the most closely approximated known samples of the two species. The same sized hiatus occurs between known localities of record for S. teapensis and the Southern Gulf enclave of S. v. variabilis. Variation elsewhere is no guarantee of intergradation or hybridization in the zone of contacts , particularly since character displacement (divergence; Mayr, 1963: 82-85) may veiy easily occur where populations of the two species actually contact. That possibility is suggested by the variation in the series of 3. v . variabilis from near Tlacotalpan and Buena Vista discussed previously; that series is from as near the range of S. teapensis as any, but shows no influence of S. teapensis. Character displacement is also evident in the number of rump rows less than 10 and number of nape rows less than 12 in populations of S. teapensis in Veracruz and Oaxaca (nearest the range of the Northern Gulf enclave of 3. v. variabilis) compared with the other populations of that species (91% vs. 80%, and 81% vs. 71%, respectively; see Table 1). On the contrary, introgression is evident in number of canthosubnasals, two occurring in 61% and 86% respectively in the two groups of S. teapensis populations. In Guatemala, conversely, these examples of displacement and introgression between S. teapensis and S. v. olloporus are just reversed. Evidence of potential sympatry there, however, suggests that these phenomena ceased there before they did in the Veracruz area, if indeed they no longer occur in the latter region. In addition, we have found two other subjective criteria that sharply distinguish the two taxa in the zone of presumed contact in southern Veracruz: the size of the lateral scales in the groin region and on sides of neck. Even the most teapensis- like S. v. variabilis seen, from 5 mi S Salinas, Veracruz (UCM 39472, with 8 rump and 1 1 nape rows), has the typical small groin scales and lateral nape scales of 3. v. variabilis (Fig. 4). These distinctions between the two taxa appear to be constant although their subjectivity makes direct comparisons frequently necessary between individuals of known identity. The relative size of the groin scales has been a particularly useful character in detecting sympatiy of the two species in two hitherto unsuspected areas of co-existence in southern parts of the range of 3, v. variabilis in Veracruz. Of two specimens from 1 km S Mocambo (just south of Cd. Veracruz), one (MZFC 218) is clearly 3. teapensis (50 mm s-v female, 47 dorsals, 9 rump rows, 1 1 nape rows, large groin scales, but canthosubnasals 3-3), the other (MZFC 220) clearly 3. v. variabilis (47 mm s-v, male, 49 dorsals, 1 1 rump rows, 13 nape rows, small groin scales, canthosubnasals 3-3). Yet a good series (UIMNH 40221-35) from 5.3 mi N Anton Lizardo, only some 20 km SE of Mocambo, are all S. v. variabilis , with small groin scales Bulletin of the Maryland Herpetological Society Page 99 Volume 29 Number 3 September 1993 and canthosubnasals all 3-3, although the dorsals vary 48-58, rump rows 9-11, and nape rows 12-14. Several series in FMNH from southwestern Veracruz, from the vicinity of different railroad stations, are especially illuminating. The entire series (1476(11)) from Achotal, near the foothills, 33 km SW Acayucan, represents S. teapensis and is fully conformant with its characteristics. In a series of 32 from Perez (1317, 1319, 1684), also near the foothills, 72 air km NW of Achotal, six represent S. teapensis (dorsals 46-48, rump rows 9(4) or 10(2), groin scales large, although two canthosubnasals occur on only two of 12 sides), all others S. v. variabilis (dorsals 48-55, rump rows 10-12, groin scales small, two canthosubnasals occur on only two of 12 sides), all others S. v. variabilis (dorsals 48-55, rump rows 10-12, groin scales small, two canthosubnasals on five of 49 sides) . Out farther on the coastal plain, at San Francisco (13 air km NNW Achotal) (1331(10)) and at Ojapa (cataloged as Otopa; 1331(9), 1312(3); 31 air km NE Achotal) all specimens are S. v. variabilis. These series amply demonstrate sympatry of these two species in this area, and together with others, reveal that S. v. variabilis extends in the coastal plain far to the southeast between the isolated uplands of Los Tuxtlas and the Sierras of Oaxaca. Near the Isthmus of Tehuantepec, however, S. teapensis apparently occupies the coastal plain to the exclusion of S. v. variabilis (Figs. 6, 7). Despite the apparent variability of interaction of the two taxa at various places where they meet in southern Veracruz, we conclude, based on irrefutable evidence of sympatiy in some areas and the infallibility of especially the character of relative size of the groin scales , that, for the present, S. teapensis is properly regarded as a full species. That conclusion requires verification, either with collections in the critical zones of contact or by field observations. As superficially similar as the two species are, visual interspecific discrimination, as a deterrent to interbreeding, seems unlikely, unless behavior is a factor. Pheromones may play a role, or interspecific isolation may depend on postmating mechanisms. Only observations in the field or in field simulations are likely to provide definitive answers. 3. Sceloporus v. variabilis, Northern Gulf Enclave. This enclave is more heterogeneous than any of the others. Variational tendencies at its southern range extremity toward character-states of S. teapensis have been noted in the preceding discussion. Toward the north, in southern central Tamaulipas, a transition occurs into the small scales and small size of S. v. marmoratus. Thus the Northern Gulf enclave is intermediate between S. v. marmoratus and S. teapensis in size of scales on the body, and S. v. marmoratus is the smallest in body size not only of these three but of the whole S. variabilis complex. Page 100 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Although in general the number of rump and nape rows is correlated closely with the size of the dorsal scales and their number from interparietal to base of tail, the correlation is not infallible. Four of seven specimens (KU 61732-8) from 10 mi NW Temapache, Veracruz, have only nine rump rows (the others have 10), but the nape rows are 13 in all four (two of the others have 14, one 13). Both high and low counts occur in the state of Hidalgo: two have nine rump rows, two have 12 nape rows, and one has 17 nape rows. Nine rump rows but 13 nape rows occur in three out of 24 from the Sierra de Juarez, Oaxaca. One of two from 15 km ESE of San Juan del la Punta, Veracruz, has 9 rump and 13 nape rows, and one specimen in 13 from 3 km SW San Marcos, Verzcruz, has a rump-nape formula of 9- 1 1 . No others with as few as 9 rump rows in the Northern Gulf enclave have been seen from north of the area adjacent to the range of S. teapensis. In that same area scattered individuals have as many as 16 or 17 and as few as 1 1 nape rows, and up to 13 rump rows. Thus the number of rump and nape rows, as well as dorsals and canthosubnasals, seems to vary independently to a certain degree throughout most of the range of the Northern Gulf enclave. The S. v. variabilis of Smith (1939) consisted solely of the Northern Gulf enclave, because the Southern Gulf enclave was not then known, and the COG enclave was referred to S. v . olloporus. The most nearly infallible objective distinction between the Northern Gulf enclave of S. v. variabilis and S. teapensis remains the dorsal scale count. Smith’s data give a 100% separation, all S. teapensis having 47 or fewer (in 103), all S. v. variabilis having 49 or more (in 197). Cole (1978) found no S. teapensis with more than 47, but found the range in S. variabilis to be 46-69. No localities were given for the latter, but they may have included areas within the range of S, v. olloporus, in which 1% of 181 have 47 or fewer dorsals, with a range of 46 to 59 (Smith, 1937); 3% have 48 or fewer, 10% 49 or fewer. Sites and Dixon (1982) made dorsal counts to the anus rather than to the level of the rear margin of the thighs held at right angles to the body, resulting in figures exceeding Smith’s (1937) and Cole’s ( 1978) by one or two scales; nevertheless no samples from the Northern Gulf enclave have counts lower than 49 except five from the southern areas near the range of S. teapensis, and on the basis of that intermediacy intergradation could be assumed. In the discussion of S. teapensis we have proposed that it actually does not occur. Only immediately to the north of the range of S. teapensis does the dorsal scale count criterion fail, and there the p re ingu Inals and lateral nuchals provide an apparently infallible criterion. Even in the area of proximity of the ranges of S. v. variabilis and S. teapensis, those two species are distinguished with at least 80% degree of accuracy by the number of rump and nape rows (see Table 1). Bulletin of the Maryland Herpetoiogical Society Page 101 Volume 29 Number 3 September 1993 4. Scelopoms v. variabilis, Southern Gulf enclave. As indicated in Table 1 , range of variation in the Southern Gulf enclave of both rump and nape rows in considerably less than that in the Northern Gulf enclave. However, only 70 specimens have been examined, from two localities, both in Tabasco. This enclave is known only from the coast of Tabasco, although since it occurs only 5 mi SW Frontera, Tabasco (UCM 39415-53), a locality only 3 1 km SW of the Campeche border, occurrence as far as the Laguna de Terminos, Campeche, is likely. On the west, the enclave is recorded from Sanchez Magallanes, only 27 km from the Veracruz border, and accordingly it probably occurs in the latter state. Somewhere, between Sanchez Magallanes and the Coatzacoalcos-Minatitlan region, its range presumably meets that of S. teapensis, but no records are available from that approximately 61 air km interval. The enclave apparently does not extend northward beyond, at most, extreme southwestern Campeche, and to the south and east it is replaced by S. teapensis , in Veracruz, Tabasco, El Peten of Guatemala, and Belize. This enclave, like others of its subspecies, is distinguishable from S. teapensis by its small preinguinals and lateral nuchals, and, more objectively, although presumably less infallibly, by its more numerous rump rows, nape rows and combined rump -nape rows; in the latter there is only a 4% and 5% overlap respectively (see Table 1). Dorsal scale counts, however, overlap extensively, as rioted in the preceding discussion, although only a small percentage of the enclave falls below 48 ( 10%). The distribution of 3 1 counts is nevertheless skewed to the lower figures, 23% falling below 49, and 35% below 50. Despite these peculiarities, the enclave is clearly referable to S. v. variabilis , not to S. teapensis, as correctly concluded by Ramirez and Gonzalez (1991). The first specimens (four) reported of the enclave were referred at that time (Cole, 1978: 8) to S. teapensis primarily because they did then appear to come from “well within the range oF that species, and had 49-51 (x 49.2) dorsals. To a considerable extent on these grounds S. teapensis was placed as a subspecies of S. variabilis by Cole, not being aware of the existence of a population of S. v< variabilis isolated north of the range of S. teapensis. 5. Sceloporus v. variabilis, COG enclave. Occurring in southeastern Oaxaca, central and southern Chiapas and adjacent Guatemala, this enclave is encircled by S. teapensis and S. smithi, and is separated by them from contacts with the other enclaves of its subspecies. Intergrades with S. v. olloporus occur in southeastern Chiapas and adjacent Guatemala (see discussion of S. v. olloporus). Its geographic range (Fig. 5) loosely interdigitates with that of S. teapensis in the vicinity of Berriozabal and Ocozocoautla, southwest of Tuxtla Gutierrez, Chiapas, without evidence of Page 102 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 intermixing (e. g. Johnson et al., 1977). Similar range approximation and possible overlap occurs near Pichucalco, in extreme northern Chiapas, with COG known 15 mi S, and typical S. teapensis 4 mi NE, of Pichucalco. Interaction of COG and S. smithi is uncertain, since specimens from critical areas of extreme southeastern Oaxaca are lacking (see discussion of S. smithi). As indicated in Table 1 , all three enclaves of S. v. variabilis are similar in all respects; the data in Sites ad Dixon on femoral pores and dorsals also agree. 6. Sceloporus v. olloporus. Sites and Dixon (1982) concluded that this subspecies is not valid, but their data, properly revised, combined with our own, support its recognition. Their data (hereinafter designated SD) show that a total femoral pore count of 2 1 or fewer characterizes 93% of the 27 counts available for Costa Rica, 9 1% of the 1 13 for Nicaragua, 85% of the 59 from Honduras, and 100% of the nine from 5-24 km NE Cd. Rio Hondo, Zacapa, Guatemala. Our data are equally supportive: 100% of 58 from Beleju, El Quiche, Guatemala (in UCM 35023-84, 44571-44606); 83% of 6 from Zacapa Dept., Guatemala (in KU 59744-6). Two discrepant sets of SD data that we here correct are (1) a tally error assigning a count of 29 to Nicaragua (which actually pertains to UMMZ 101212, S. v. variabilis from 2 km N Miramar, Veracruz (south of Tampico, Tamaulipas), for which 24 was otherwise the highest); and (2) a figure of only 60% 21 or fewer for 10 specimens from El Salvador. We did not examine the SD specimens from El Salvador, but 92% of our counts (12) from El Salvador material (in KU 184254-69) are less than 22 (one exception, of 22); it is very difficult to make accurate counts of femoral pores on most females and juveniles, and we reject the SD counts as inaccurate in this particular series, in view of our different results obtained with great care. These figures are consistent over a wide area from Costa Rica into Guatemala, and are in sharp contrast with the SD data for S. v. variabilis from Mexico. Excluding their samples of S. teapensis and S. smithi, the following percentages obtain of total femoral pore counts less than 22, arranged more or less in N-S order. The Gomez Farias region, Tamaulipas (SD locality 5), 4% of 25; 2 km N Miramar, Veracruz (south of Tampico), 3% of 31; Valles region, San Luis Potosi (SD locality 7), 0% of 20; Queretaro (SD locality 8), 0% of 235 (total range is 22-34, not 14-34 as in SD; the single 14 count is a lapsus for 14 + 17 (total 31) fide recheck by Dixon); Tecolutla region, Veracruz (SD locality 9), 0% of 16; Xalapa, Veracruz, 0% of 9; Cd. Veracruz, 3% of 41; 19 km N Alvarado, Veracruz, 10% of 10; 8 km N Lerdo de Tejada, 0% of 14; Cosolapa, Oaxaca, 0% of 26; Soyaltepec.Tuxtepec dist., Oaxaca, 9% of 1 1; Sanchez Magallanes, Tabasco, 8% of 12; Playa Miramar, Bulletin of the Maryland Herpetological Society Page 103 Volume 29 Number 3 September 1993 1 1 km SW Frontera, Tabasco, 4% of 24; Chiapa de Corzo, Chiapas, 1 1% of 9; Tonala, Chiapas, 8% of 12; and Jacaltenango, Guatemala, 8% of 25. The data for the Playa Miramar series are our own, based on UCM 39415-53, of which only 24 could be counted reliably; the 1 5 SD counts, based on UIMNH material taken with the UCM series, and of which 53% are lower than 22, we are convinced are inaccurate, since most specimens involved are females or juveniles, which are usually very difficult or impossible to count accurately. Thus throughout the extensive ranges of both S. v. variabilis and S. v. olloporus , except for a small intervening area in southwestern Guatemala and adjacent Chiapas, the two subspecies are consistently different, without evident clinal convergence . The level of conformance with the stated criterion is at least 89% in any given area represented by the 520 tallied S. v. variabilis (an over- all conformance of 98%, 12 exceptions), and is at least 83% in any given area represented by the 275 tallied S. v. olloporus (an over- all conformance of 92%, 22 exceptions). Excluded from the tallies for S. v. olloporus is a series of 2 1 specimens (in UTA 8948-9, 20329-32; KU 59717-30, 157471-2, 187174-8, 187180-2) from El Progreso Dept., Guatemala, which have only 67% with less than 22 femoral pores; part of that series was included in the SD data (locality 32), with a comparably low percentage (73%). The lowest percentage elsewhere is 83% for Zacapa Dept. , Guatemala. Only in this area of Guatemala is there clear evidence of an approach of S. v. olloporus to the characteristic numerous femoral pores of S. v. variabilis . Yet typical populations of the latter subspecies are nearly 150 km to the west, with typical populations of S. v. olloporus intervening. It seems likely that their ranges were more nearly approximated in the past, and that the El Progreso populations reflect that approximation to some extent. Unfortunately sufficiently large series from sufficiently numerous localities in central and western Guatemala are not available as yet to provide a satisfactory picture of distribution of the subspecies there, and their areas of intergradation. However, at least four series referable to S. v. variabilis show the influence of S. v. olloporus, all from the Chiapas-Guatemala border south of 16° north latitude, and all (SD data) with 33-40% of the femoral pore counts less than 22. In Chiapas. 40 from Comitan have 40% (16) with fewer than 22 femoral pores, and 14 from Zapaluta have 37% (7); in Huehuetenango, Guatemala, 9 from Rio Cuilco have 33% (3), 12 from 2 km SW Colotenango also have 33% (4), and three from 33 km WNW Huehuetenango, El Tapon area (our data) have 33% (1). Page 104 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Stuart (1954) long ago deciphered the distribution of these two subspecies in Guatemala, mapped their locality records, noted apparent intergradation exactly where indicated here (at Cambal, on Rio Cuilco), and declared that neither is particularly closely related to either S. cozumelae (lapsus for S. chrysostictus) or S. teapensis , both of which occur nearby on the Caribbean versant. The most conspicuous geographic variation observed in S. v< olloporus is in number of postrostrals, 70% (in 62) of which are less than 4 in Costa Rica material, 68% (in 196) in Nicaragua specimens. In Honduras material (22 specimens), 32% have fewer than 4, much as in S. v. marmoratus (33% in 44), and in El Salvador 23% (3 of 16) have fewer than 4. All other samples of S. v. olloporus, S. v. variabilis , S. smithiand S. teapensis have no more than 19%, and as little as 6%, with fewer than 4 postrostrals. Unless other distinctions are found in the Costa Rica-Nicaragua populations, no taxonomic separation of them from S. v. olloporus seems justified, although their postrostral deviation from other populations suggests that genetic exchanges with more northwestern populations may be reduced. 7. Sceloporus v. marmoratus . This diminutive subspecies, the northernmost of the complex, extending into Coahuila on the northwest (Gonzalez et al., 1989), poses no major problem taxonomically. Its small size and numerous dorsals are its most distinctive characteristics. Counting the dorsals even to rear margins of the thighs, Smith (1939) found 91% of S. v. marmoratus with 60 or more, 93% of S. v. variabilis with 59 or fewer. However, the data taken by Sites (for Sites and Dixon, 1982), with dorsal counts taken to the level of the anus, do not provide such a sharp separation. Their locality areas 1-4, in Texas and central Tamaulipas, yield the best separation with 79% (of 173) having 61 or more, as compared with only 26% (of 90) in their areas 5, 6, and 7 in southern Tamaulipas, southern San Luis Potosi and extreme northern Veracruz. The separation is better for their locality areas 9 and 10 incentral Veracruz (17% 61 or more in 24 specimens), their locality area 8, in Queretaro, is not considered here because an upland race, probably extending into Hidalgo (Mendoza, 1990) maybe involved that has not yet been studied carefully. Maximum size conforms with the geographic parameters dictated by the dorsal scale counts. In 174 specimens measured by Sites (in Sites and Dixon, 1982) from their locality areas 1-4, only one reached 57 mm s-v (measuring only 57.4 mm), and only 28 (16%) measured as much as 50 mm. Their 91 measurements for locality areas 5-7, on the contrary, reached a maximum of 71.9 mm, and 66 (73%) measured 50 mm or more. That large size is maintained throughout the range of S. v . variabilis and S. v. olloporus , except perhaps in the Tabasco series, which averages smaller than other Bulletin of the Maryland Herpetological Society Page 105 Volume 29 Number 3 September 1993 samples in s-v length. The maximum in that series of 70 is 67 mm, and only 43% measured 50 mm or more - figures still distinctly greater than those of S. v. marmoratus. The small size of the dorsal scales is reflected not only by the standard dorsal scale count but also by the numerous nape and rump rows (84% 15 or more, 87% 12 or more, respectively). Only S. smithi of the S. variabilis complex is similar, with 50% and 85% respectively. All other populations (see Table 1) have 18 or 19% or less, respectively, except for the Northern Gulf enclave of S. v. variabilis , with over-all percentages of 29 and 19, respectively and slightly higher percentages to the north (32 and 22, respectively). S. v. marmoratus and S. smithi likewise have the highest summed nape and rump counts, and of those totaled with the canthosubnasal count (see Table 1). Despite the small size of its dorsal scales, S. v . marmoratus lacks the lateral ridge fold characteristic of S. smithi the only other taxon now recognized with small dorsals. Curiously, S. v. marmoratus has the highest frequency (23%) of three or fewer postrostrals outside of southern (Nicaragua, Costa Rica) S. u. olloporus (68%); northern S. v. olloporus is about the same (22%) (see Table 1). Equally strange is the relatively high frequency of only two canthosubnasals (31%) in S. v. marmoratus , a feature usually characteristic of S. teapensis and occurring elsewhere to no more than 10% of the specimens examined, except for the COG enclave of S. v. variabilis , with 21% (see Table 1). The range of S. v. marmoratus in Texas is thoroughly detailed in Axtell (1988); comparable detail in Mexico awaits the future, but it is clear that the area of intergradation with S. v. variabilis lies very close to or blankets the Tropic of Cancer in at least part of its extent. It lies close to El Mante, Tamaulipas; three specimens (KU 61743-5) from 29 mi E El Mante, Rio Guayalejo, are clearly representative of S. v. marmoratus (13 rump rows, 16- 17 nape rows, 56 mm s-v- maximum), whereas 5 mi WEI Mante (KU 61740- 1) the sample clearly represents S. v. variabilis ( 1 1 rump rows, 15 nape rows, 69 mm s-v max.), as does another specimen 12 mi S El Mante (KU 61739, with 10 rump and 13 nape rows). Apparently S. v. marmoratus extends farther south near the coast, and S. v. variabilis farther north near the Sierra Madre Oriental. Page 106 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Acknowledgments We are much indebted to the authorities of the museums from which specimens were borrowed for study: Dr. Daniel B. Blake (UIMNH), who also kindly researched locality data for us; Dr. Jonathan A. Campbell (UTA), who also provided much help with data on specimens and localities in Mexico and Guatemala; Dr. Darrel R. Frost (AMNH), who also kindly checked erroneous identifications and proffered taxonomic counsel; Alan Resetar (FMNH); Jose Rosado (MCZ), who also traced records in an attempt to resolve the problematic “S. smithi x S. v. variabilis intergrade”; Greg Schneider (UMMZ), who also valiantly tried to shed light on the same intergrade problem as well as others; Dr. Philip J. Regal (JFBM); John Simmons (KU), who also searched records to eliminate discrepancies and ambiguities of locality citations; and Dr, Shi Kuei Wu (UCM) , who also provided facilities for study. In addition we are very grateful to Dr. Jack W. Sites, Jr., for the use of his extensive data on the S. variabilis complex; and to Dr. William B. Lewis for space, facilities and financial support for the project. Fernando Mendoza Quijano kindly permitted us to examine, list and plot localities on our map of MZFC specimens he recently collected. Key to Members of the Sceloporus variabilis Complex In the following key, the dorsal scale count is interpreted as the least count between interparietal and the level of the rear margin of the thighs held at right angles to the base of tail. The rump row count is the least dorsal scale count across the sacral region anterior to the level of midthigh but posterior to the level of anterior margin of the thighs held at right angles to the body axis. The nape row count is the least dorsal scale count across neck between the levels of arm insertion and ear openings. 1 A. Dorsal scales 49 or fewer (about 98%); rump rows 7-9 (85%; no other taxa more than 3%, although variation in 5 regions of variabilis is 0- 18%); rump, nape and canthosubnasal total count 20-27 (97%; no other taxa more than 6.2%, although variation in 5 regions of variabilis is 3- 13%); scales just anterior to groin relatively large, more than half size of median lateral abdominals; median lateral nape scales about same size as enlarged scales on crest of lateral nuchal fold . . . . . . . . . . teapensis Bulletin of the Maryland Herpetological Society Page 107 Volume 29 Number 3 September 1993 B. Dorsal scales 50 or more (about 98%); rump rows usually 10 or more (all taxa no less than 97%, although variation in 5 regions of variabilis is 82-100%); rump, nape and canthosubnasal total count 28 or more (all taxa no less than 93%, although variation in 5 regions of variabilis is 87-97%); scales just anterior to groin relatively small, much less than half size of median abdominal laterals; median lateral nape scales distinctly smaller than enlarged scales on crest of lateral nuchal fold.. . . . . . . . . . . . . . . . . . 2 2 A. Ground color uniformly very dark, with no or faint dark spots evident on sides or between prominent dorsolateral white stripes; latter usually 2 and 2 half scale rows wide at widest part; adult females with abdominal semeions well developed; hatchlings pink- tailed; rump rows 12 or more (85%); keels on dorsal body scales forming conspicuous, continuous fine ridges; median lateral abdominal scales little larger than border axillary and groin scales, lying on a longitudinal ridge; to 78 mm s-v . . . . . . smithi B. Ground color gray to brown, with dark spots readily evident on sides of body and in two paravertebral series median to dorsolateral light stripes; latter not so prominent or wide; abdominal semeions not or but dimly evident in adult females; hatchling tail of same ground color as body; rump rows variable; keels on dorsal body scales not forming conspicuous, continuous ridges; median lateral abdominal scales much larger than border axillary and groin scales, not on a longitudinal ridge; maximum s-v length 76.4 mm . . . 3 3 A, Dorsal scales 60 or more (91%); rump rows 12 or more (87%); maximum s-v length 57.4 mm . . v. marmoratus B. Dorsal scales 59 or fewer (93%); rump rows 1 1 or fewer (83% for all enclaves, which vaiy 78-100%); maximum s-v length 76.4 mm ....4 4 A. Femoral pores total 21 or fewer . . . . . . v. olloporus B. Femoral pores total 22 or more . u. variabilis Page 108 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Literature Cited Axtell, Ralph W. 1988. Sceloporus variabilis . Interpretive Atlas of Texas Lizards (4): 1-11, map 3. Bocourt, Marie-Firm in. 1874. Deux notes sur quelques sauriens de lAmerique tropicale. Annls. Sci. Nat. Paris, (5) 19(4): 1-4. Boulenger, George Albert. 1 897. A revision of the lizards of the genus Sceloporus. Proc. Zool. Soc. London, 1897: 474-522, pi. 33 (color). Campbell, Jonathan A. and Jay P. Vannini, 1989. Distribution of amphibians and reptiles in Guatemala and Belize. Proc. Western Foundation Vert. Zool. ,4(1): 1-21, figs. 1-23. Cole, Charles J. 1978. Karyotypes and systematics of the lizards in the variabilis , jalapae and scalaris species groups of the genus Sceloporus. Am. Mus. Nov., (2653): 1-13, figs. 1-4. Gonzalez, A., Arturo, Fernando Mendoza Q., Richard R. Montanucci and Ernest A. Liner. 1989. Una colecta herpetologica en el norte de Coahuila. VII Coloquio de Investigacion, Esc. Nac. Estud. Profesionales Iztacala-UNAM: 26. Guillette, Louis J., Jr., John Weigel and Gary Flater. 1983. Unilateral testicular pigmentation in the Mexican lizard Sceloporus variabilis . Copeia, 1983(1): 155-161, figs. 1-4. Gunther, Albert C. L. G. 1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. London, Porter, xx, 326 pp., 76 pis. Hallowell, Edward. 1852. Description of new species of reptiles inhabiting North America. Proc. Acad. Nat. Sci. Philadelphia, 6: 177-182. Bulletin of the Maryland Herpetological Society Page 109 Volume 29 Number 3 September 1993 Hartweg, Norman and James A. Oliver. 1937. A contribution to the herpetology of the Isthmus of Tehuantepec. I. The scelopori of the Pacific slope. Occ. Pap. Mus. Zool. Univ. Michigan, (356): 1-9. Johnson, Jerry D., Charles A. Ely and Robert G. Webb. 1977. Biogeographical and taxonomic notes on some herpetozoa from the northern highlands of Chiapas, Mexico. Trans. Kansas Acad. Sci., 79(3, 4): 131-139, figs. 1-2 (1976). Lee, Julian C. 1980. An ecogeographic analysis of the herpet ofauna of the Yucatan Peninsula. Univ. Kansas Mus. Nat. Hist., Misc. Publ., (67): 1-48, 22 figs., 27 pis. Levi ton, Alan E. et al. 1980. Museum acronyms - second edition. Herp. Rev., 11(4): 93- 102. Mather, Charles M. and Jack W. Sites, Jr. 1985. Sceloporus variabilis Wiegmann. Cat. Am. Amph. Rept., (373): 1-3, map. Mayr, Ernst. 1963. Animal species and evolution. Cambridge, Mass., Harvard Univ. Press, xv, 797 pp., ill. Mendoza Quijano, Fernando. 1990. Estudio herpetofaunistico en el transecto Zacualtipan- Zoquizoquipan-San Juan Meztitlan, Hidalgo. Tlalnepantla, Mexico, Biol. Dissert, vi, 97 pp. Meyer, John R. and Larry David Wilson. 1973. A distributional checklist of the turtles, crocodilians, and lizards of Honduras. Los Angeles Co. Nat. Hist. Mus., Contr. Sci., (244): 1-39. Ramirez-Bautista, Aurelia and Alberto Gonzalez- Romero. 1991. Notes on the reproduction of the Rosebelly lizard, Sceloporus variabilis (Sauria; Iguanidae) from Dos Bocas, Tabasco, Mexico. Bull. Chicago Herp. Soc., 26(12): 270-272, figs. 1-2. Sites, Jack W., Jr. and James R. Dixon. 1982. Geographic variation in Scleoporus variabilis, and its relationship to S. teapensis (Sauria: Iguanidae). Copeia, 1982(1): 14-27, figs. 1-6 . Page 1 10 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 Smith, Hobart M, 1937. A synopsis of the variabUis group of the lizard genus Sceloporus » with descriptions of new subspecies. Occ. Pap. Mus. Zool Univ. Michigan, (358); 1-14. 1939. The Mexican and Central American lizards of the genus Sceloporus. Zool. Ser. Field Mus. Nat. Hist., 26: 1-397, figs. 1-59, pis. 1-31. _____ , Matthew S. Rand, J. David Drew, Bruce D. Smith, David Chiszar and Christine M. Dwyer, 1991. Relictual intergrades between the Northern prairie lizard (Sceloporus undulatus garmani ) and the Red-lipped plateau lizard (S. u. erythrocheilus) in Colorado. NW Naturalist, 72: 1-11, figs. 1-8. Stuart, Laurence C. 1943. Comments on the herpetofauna of the Sierra de los Cuchumatanes of Guatemala. Occ. Pap. Mus. Zool. Univ. Michigan, (471): 1-28, figs. 1-7, pi. 1. 1954. A description of a subhumid corridor across northern Central America, with comments on its herpetological indicators. Cont. Lab. Vert. Zool., Univ. Michigan, (65): 1- 39, maps 1-6, pis. 1-6. 1971. Comments on the malachite Sceloporus (Reptilia; Sa uria: Iguanidae) of southern Mexico and Guatemala. Herpetologica, 27)3): 235-258. Wiegmann, Arend F. A. 1 834. Herpetologica mexicana seu descriptio amphibiorum Novae hispaniae. Pars prima. Saurorum species. Berlin, Luderlitz. vi, 54 pp., 10 pis. Appendix Specimens we have examined, or that have been examined for us, in the course of this study are as follows. Sceloporus smithi OAXACA . AMNH: MLabo Sepas\ nr Tehuantepec (66897). KU: 2 mi BSE Tehuantepec (42049); 3 mi NW Tehuantepec (37806); Bulletin of the Maryland Herpetological Society Page 1 1 1 Volume 29 Number 3 September 1993 6 mi WNW Tehuantepec (42045-8). UCM: JuchitanDist.: Cerro de Potosi, nr Chahuites (44425-33); Palomares (39852); Tehuantepec Dist.: Cerro San Pedro (39856); El Limon (39855, 41131-5); Escurano (39853-4); Tamasola, nr Magdalena Tequisixtlan (39857-9); Tlacolula Dist : 2 mi S Totolapan, 3700’ (16698); YautepecDist; San Juan Acaltepec (44424). UIMNH: Yautepec Dist.: 1.9 mi SE La Reforma (40177). UMMZ: Yautepec Dist.: 5.4 mi NW El Camaron, 2100' (114942); 7.9 mi SE El Camaron, 4100' (126227). Sceloporus smithi x S. teapensis hybrids (referred to S. smith i). OAXACA. KU: Yautepec Dist.: 2 mi N, 6 mi W Nejapa (43733-4). Sceloporus teapensis. BELIZE. KU; Cayo: 1.1 mi S Georgeville (157451); 31.5 mi S Georgeville (157448-9); Rio (157450); Xunantunich (144946, 171 505-7) ; Stann Creek: 13.2 mi SW Silk Grass (1 57452). CHIAPAS. KU; Palenque ruins (94096-9); 3.5 km NW Pichucalco (94102); 4 mi NE Pichucalco (94100-1). UCM; Palenque (49668-72, 49674). GUATEMALA. KU; AltaVerapaz: Finca Chicoyou, nr Coban, 980 m (59715-6); ElPeten: 2.5 mi W El Cruce (157460-1); 34.6 mi E El Cruce (157468-9); 48.2 km W Melchor de Mencos (171482); 4 mi N Poptun (157453-5); 9.9 mi NW Poptun (157456); 12. 1 mi NW Poptun (157457-9); 2.5 km N Ramate (171488-504); 8.3 mi SW San Benito (171483-7); 3 mi S Tikal (157463-7); 14.2 mi S Tikal (157462); 15 km SE La Libertad, Toocog (55808-9, 55811, 55816-24, 55826-34, 59712); Izahal: Las Dantas, WEI Estor, 30 m (187173). UCM; El Peten: Sayaxche (22275). UTA; Alta Verapaz: 5. 1 mi NE Coban (33618-20); El Quiche: Finca La Perla, El Tesoro (20333-5, 20748-51); Izahal: El Estor, El Coupon (29258); El Estor, El Zapotillo (29279); Sierra de Santa Cruz (E side Cerro 1019,byAldeaLa Libertad, 29280, 24294-6; Cerro Cana Tomasa, S side, 29259-66, 29278, 33624-7; S side Cerro La Dicha, 29285; Chichipate, 23575-6, 29281-4; Chinamococh, 29267-9; Finca Semuc, 3rd km Shdqtrs, 29838; Finca Semuc, 4.5 rd km S hdqtrs, 22110-1; Marc aj am, 29270-7; Seshan, 23577-85, 29286-93); ElPeten: San Jose (33621-3); San Francisco (23586); Tikal (22109). OAXACA. AMNH; Ocotal, betw Matias Romero and Santa Maria Chimalapa (65029-30). FMNH: Santiago Guevea (113898- 113903). KU; 20 km S Jesus Carranza, Veracruz (24260, 24382, 24384); Tollocito (39684-5); 2 mi ETollocito (44857); 2 mi STollocito (44858, 44860- 3, 44873-4); 2 mi STollosa (33796). MZFC; Jalahui, 290 m (5421); 1.47 km W Jalahui (5422-4). UCM; 12 de Julio, Donaji (44423); Palomares (39849- 51); Real de Sarabia, 7 mi S (18984); Tamasola, Tequisixtlan (39848). UIMNH; Coatlan (37343). VERACRUZ. FMNH; Ochotal (1476 (11)); Perez (1317(3), 1319(3)). JFBM: nr Cerro Balzapote (10178, -87, 10201, -3, -14, -62-8, -70, -89, 10469-70). KU; 5 mi S Catemaco (39686, 44859); 20 km ENE Jesus Carranza (27519-23); 3 km E San Andrds Tuxtla (24162,24249, -67, -9, -71, -5, 24379). MZFC; 6 km NE Coyame (FMQ 2178-9); E slope Volcan Bulletin of the Maryland Herpetoiogicaf Society Page 112 Volume 29 Number 3 September 1993 San Martin (FMQ 2156-4); 1 km S Mocambo (218). UCM: Coyame, Lake Catemaco (28773-7, 28779-81, 28783-28809, 28811-45); 4 km N Coyame (28846-64). Sceloporus v. marmoratus . NUEVO LEON. KU: 4 mi W Allende, Rio Ramos (68106, 68108); Parajede losOsos, Santiago (92608-1 1). TAMAUUPAS. KU; 8 mi S Llera (61747); 20 mi E Mante, Rio Guayalejo (61743-5). TEXAS. KU: Atascosa Co. (Benton, 7227); Bexar Co. (Helotes, 1 1005-8); El Paso Co. (El Paso, 15572; locality erroneous, Axtell, 1988: 6); Frio Co. (Dilley, 12468; llmiWDilley, 15192-8); Lasalle Co. (15 mi NEncinal, 15205-6); Nueces Co. (15 mi SE Corpus Christi, 88344-9); San Patricio Co. (7 mi NE Sinton, 61729-31); Starr Co. (Rio Grande City, 15354-5; Los Olmos Bridge, Rio Grande City, 15199-203; 3 mi NE Rio Grande City, Hy755, 145800; 6 mi W Rio Grande City, Arroyo El Salado, 12467); Webb Co. (17.3 mi S jet Hy 59 & 2050, 145801-3). Sceloporus variabilis olloporus. COSTARICA. KU: Guanacaste : 10 km WBagaces (102442-4); Comelco (157792); El Coco (67296; 2 km SW, Ojotal, 67290-5; Playade, 100709-41, 125552-60); Guardia( 3 kmN, Rio Tempisque, 125551; 10 kmN, 102445); La Cruz (27 km S, 212 m, 95519; 44 km S, 133 m, 95520); betw Los Angeles and Tilaran (36861); 23 km S Pena Blanca, 227 m (95518); Sardinal (125561); Puntarenas (34846): Maribella Hotel (34226- 44). EL SALVADOR. KU: Cuscatlan : Tenancingo (Rio Tizapa, 184255, 184257-61; 2.8kmNE, Rio Quezalapa, 184264-6); San Miguel: San Antonio Chavez, 10 km NNE San Miguel (184254, 184256-7); S Side Volcan San Miguel (184262-3); Santa Ana: Metapan (6kmS, 184268; 7km SE, 184269). GUATEMALA. KU: Baja Verapaz : 18 km N Salama, 1500 m (59744-6); Zacapa: Gualan, Aldea Doha Maria, 200-450 m (190787-9); Rio Hondo (5 km SW, 225 m, 59732-43; 24 km NE, 150 m, 59731); Sierra de las Minas, 6.4 km N jet CA-9 and rd to San Lorenzo, 500 m (190784-6); 6 km E Zacapa turnoff from CA 9, Rio Piedra de Filar, 171 m (187179). UCM: El Quiche : Baleju, 10 air km SW San Cristobal Verapaz, Alta Verapaz (35023-84, 44571-44606). UMMZ: El Quiche : Sacapulas, 1225-1275 m (120180(3)); Zacapa: Finca San Jorge, 3 km NE Uzumatlan, 600 ft (107072). UTA: Zacapa: Cabanas, Aldea El Rosario (29839). HONDURAS. KU: Choluteca: Choluteca, Agua Caliente de Pavana, El Pital (192322); Comayagua: 13.7 km E Siguatepeque, 1500 m (67286-9); El Paraiso: Guinope, 4.2 km NW, 1435 m (209318-9); 30.6 km NW Mandasta, 1500 m (209320-2); Soledad, Aldea las Marias, Los Canales (192323-4); Intibuca: 15 km E La Esperanza, 1490 m (194331-3); Lempira: betw Gracias and Villa Verde, 1030 m (200579); 3 km N Gualcince, 1510 m (194330); Olancho: 10.5 km S San Esteban, 470 m (200577); Valle: Isla Zacate Grande (194334); Yoro: 2 km S Coyoles, Rio Aguan (101442-3); Montana de Ruidosa, above Calpules, 1000 m (200578). NICARAGUA. KU: Boaco: 3 km E Tuestepe, 160 m (103266); Bulletin of the Maryland Herpetological Society Page 113 Volume 29 Number 3 September 1993 Carazo : 3 km N, 4 km W Diriamba, 600 m (113006); 1 mi SE Masachapa (42036-44); Chinandega : Hda. Bellavista, Volcan Casita, 720 m (101872); 4 km N, 2 km W Chichigalpa (85886-94); San Antonio, 15 m (85896); foothills N slope Volcan San Cristobal (85963-4). Chontales : river 8 km W Muhan (174090-1); 1 km N, 2.5 km W Villa Somoza, 330 m (1 13004-5); Estelt Esteli ( 8 km N, 770 m, 67297; 8 mi NNW, 42013; 10 km N, 740 m, 1 16972-3); Finca Daraili, 5kmN, 14 km E Condega, 940 m (85874-84); Granada: Finca Santa Cecilia, 6.5 km SE Guanacaste, 660 m (101871); Volcan Mombacho (174092); Jinotega: nr Apanas (174088-9); Leon: Ingenio San Antonio (174082); Poneloya (174080-1); Managua: Cerro Coyotepec (174084-5); 23 km N Las Maderas, 394 m (95521-2); Managua (3 mi SW, 42027-35; 6 mi WSW, 42026; 15 km S, 174083); nr Masachapa, 182 m (95523); Sabana Grande (2 km N, 50-55 m, 84869, 85863-8, 85873; 2 mi N, 42 120-5 1 ; 3 km N, 50 m, 85869-72); 4 mi E San Lorenzo (42017-25); Tipitapa (174086-7; 1 kmS, 50 m, 85885); Matagalpa: 1 1 mi SE Dario (42014-6); 3 km NWSebaco, 440 m (103267); Rivas: Finca Arnayo, 13kmS, 14 km E Rivas, 40 m (85897- 85927); 1-4 km NNE Moyogalpa, Isla Ometepe, 40 m (85895, 85950-62, 85965-9); Rio Javillo, 3 kmN, 4 km WSapoa, 40m (85928-30); 4.5 km E San Jose del Sur, Isla Ometepe, 60 m (85931-49); Zelaya: 3-4 km NNW Puerto Cabezas, rd to Waspan, 30 m (101444). Sceloporus v. olloporus x S. v. variabilis (referred to S. v. olloporus ). GUATEMALA. KU: El Progreso: El Rancho (2 km N, 275 m, 59717-30; N bank Rio Motagua across from, 317 m, 18174-7); 27.8 mi SWRio Hondo, Zacapa (157471-2); jet Rio Morazan and CA-14, 329 m (187180-2); jet Rio Huyus and CA-9, 268 m (187178). UTA: El Progreso: El Rancho (7.3 mi W, 8948-9; across Rio Motagua from, 20329); Rio Morazan, 10.6 km Wjct CA-9 and CA- 14 (20330-2). Sceloporus v. variabilis (Northern Gulf Enclave). HIDALGO. JFBM: 7.7 mi S Chapulhuacan (2865). KB: 6 mi NE Jacala (61750); 4 km S Tehuatlan (24255); 5 km S Tehuatlan (23819, 24152, -7, -60, -68, -71, 24253-4, -62, 24381, -6); Rio Chinameca, 7.2 km NW Tianguistengo (54056). OAXACA. KU: Vista Hermosa (87395, 87482). UTA: Metates, N slope Sierra de Juarez (11877-82, 24022-39); 1 mi S Valle Nacional (8451-4, 8480). PUEBLA. JFBM: Mesa de San Diego, Rancho El Alengihre (2238). SAN LUIS POTOSL JFBM: Xilitla rd, 3.3 mi SW Hy 85 (2691, 2694). KU: 1 1 km N Antiguo Morelos (23310); El Salto Falls (61748, 95525); 1.5 km N Rio Frio (95524); 3 mi NE Tamazunchale (61749); Xilitla region (27049-55). UCM: 3.9 mi NE Cd. del Maiz (48963-76); El Salto (28755-65, 28767-72). TAMA UUP AS. JFBM: Rancho del Cielo, Gomez Farias (2233). KU: 1 mi S Altamira (33987- 8); 5 mi W Mante (6 1 740- 1 ) ; 1 2 mi S Mante (6 1 739, 6 1 742) ; 24 km N Rio Frio (95526); 2 mi W Tampico (61746). VERACRUZ. FMNH: “Otopa” (= Ojapa) (1311(9), 1312(3)); Perez (1317(5); 1319(11); 1684(10)); San Francisco Page 1 14 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 (1331(10)). JFBM: 1.8 mi S Buena Vista (9.3 mi N Tlacotalpan), 2852, -8, 2860-1); 2 mi S Tlacotalpan (2865). KU: Alvarado (26736-7, -40, -42-3, -52, -54, -56, -61, -64-6, 27037, 44383); 4 km S Alvarado (59706-10); Boca del Rio (26747, 27042); 5 km SW Boca del Rio (24338-9, 24348-9, 24351); 13 ml ESE Boca del Rio (39687); betw Boca del Rio and Veracruz (156260-1); Cerro Gordo, 7 km NNW (23822, 24172, 24250, -70, -72, -77, 24380); Coscomatepec (26738, -63); Cuautlapan (105817, -19-20); 5 km ENE El Jobo (24268); 5 km N Jalapa, 4500 ft (24333-7, -42-4, -50, -52-3); 17 km SE LasTrancas, Hy 140 (95527-31); Mirador, 3500 ft (23823, 24261, 24385); Mocambo Beach (158788); 9 km NW Nautla (24164, 24265, -74); 4 km W Paso de San Juan (24159, 24251, -7, -76); 20 km WNW Piedras Negras, Rio Blanco (23256, -62-3, -5-7, 23309, -11. 24256); 3 km W Plan del Rio (24346); 5 km W Plan del Rio (24340-1); 5 km S Potrero, 1700 ft (26739, -50, -7); 8 km S Potrero, Sala de Agua, El Maguey (23308); 13 km WNW Potrero, 2000 ft (26748, -51); Potrero Llano, 350 ft (27040); Potrero Viejo, 1700 ft (24153, -74, 24378, 26735, -41, -60); Puente Nacional (24252, 27034, -9); 8 km NW Rinconada, Hy 140 (95532); 15 km ESE San Juan de la Punta (24345, -7); 3 km SW San Marcos (23820-1, 24156, -8, -63, -5, -7, -9-70, 24263, -6, -73. 24389); 10 mi WNW Temapache (61732-8); Teocelo (26749, -55, -8-9, -62, 27035-6,-8); 1 5 km ENE Tlacotepec, 1500 ft. (24166, 24258, 24387); 4 km W Tlapacoyan, 1700 m (24155, -61, -73, 24259, 24388). MZFC: Punta Arenas, 6 km NLerdodeTejada (FMQ 2141-3); 1 km S Mocambo (220). UCM; 1.1 mi SW Acutlzingo (48393-4); 15 mi N Alvarado (50767); El Morro Lighthouse (39455-9); La Palma, Mpio. Totutla, 1200 m (30350-2); 1 mi N Punta del Morro (38365-72); 18.6 mi W Rinconada (48395-9); beach 5 mi S Salinas (39460-72); 4 mi S Tecolutla Ferry (39454); coast rd 20 mi SE Veracruz (45615). UIMNH: 5.3 mi N Anton Lizardo (40221-35). Scelopoms v. variabilis (Southern Gulf Enclave). TABASCO. MZFC: Punta Sur de Sanchez Magallanes (FMQ 2198-2205); Ejido El Alacran, 20 km ESE Sanchez Magallanes (FMQ 2206). UCM: 5 mi W Frontera, Playa Miramar (39415-53). UIMNH: 1 1 km SW Frontera, Playa Miramar (87382- 99); Sanchez Magallanes (87368-81). Sceloporus v. variabilis (Chiapas, Oaxaca, Guatemala Enclave). CHIAPAS. KU: Rio Jesus, 32 km NW Pijijiapan (67283); Linda Vista, 2 km NW Pueblo Nuevo Solistahuacan (59747); 12.8 km ESETeopisca (187187- 8). MZFC: 10 km NE Frontera deComalapa (FMQ 2235-7). UCM: Mt. Ovando (39673-4); 6.4 mi N Tuxtla Gutierrez (48954-8); 15.3 mi N Tuxtla Gutierrez (48959-61). UIMNH: 15 mi S Pichucalco (8898-9); 13 mi S La Trinitaria (8997). UMMZ: Arriaga (88404); Bochil (99850); 35-36.2 mi S Jitotol (119850(3), 119851(2), 119852(4)). UTA: 11.3 km ESETeopisca, 2073 m (33632); Teopisca (6.4 mi ESE, 6088-93; 10.3 km ESE, 5762-3; 10.5 km ESE, 11862; 11.3 km ESE, 11863-7); Tulanca, 12.1 km ESE Teopisca Bulletin of the Maryland Herpetological Society Page 1 15 Volume 29 Number 3 September 1993 (11872-4); Tulanca quarry (24040-1); 1 mi W Tulanca (8929-31); 1.1 mi E Tulanca (8927-8, 8950). OAXACA. AMNH: Cerro Atravesado, 4000ft (66894); Rio Chicapa, nr Cerro Atravesado, 1800 ft (66939). FMNH: Niltepec (1468, 99251-2). MCZ: Tapanatepec (33454-5). UIMNH: Niltepec (21954); Plan de Campana, Rio Porto Moneda (37256, 37264). UTA: Cerro Haul (5674). GUATEMALA. KU: Huehuetenango: El Tapon, 1390 m, 33 km WNW Huehuetenango (116960-2). Scelopoms v. variabilisx S. v. olloporus (referred to S. v. variabilis). CHIAPAS. KU: 22.8 mi SE Comitan, Hy 190 (145231); Las Margaritas (94103). UCM: 15.8 mi SE Comitan (48950); 22.8 mi SE Comitan (48951- 3); 41 mi S Comitan (18985). GUATEMALA. KU: Totonicapan: Santa Lucia La Reforma, 1700m(lkmE, 190793; 4 km N, Sacsiguan, 190791-2; 8 km NW, Rio Sajcoclaj, 190790). Page 1 16 Bulletin of the Maryland Herpetological Society Volume 29 Number 3 September 1993 CO z i o > (0 > . . Q. « ,® ® d x: > m o ® 1 ® Q s s a 3 o * * E CO * ® 2 ^ i r0 o Z. .53 > « «> a. iff I si • ^ o VJ o w K 5 jg t °1 ■> a (0 CO* o (J ^2 d 2^ % - W 7J ■ ,9 ® c «* iS <0 O ~ ^ D C 3 o> ® O <0 _ to 3 E M o ® • z^£ - E 3 i <3 ® o g C , 3 & > ^1 = ! £ •§ « i . 2 ®" ;| ^ J ! ^ co 1 L« Z =3 imgo : | « c » S « o : 3 E 2 ‘ O 2 .w 13 0® ! Is 2 ' ^ 2 > r ^ O co Bulletin of the Maryland Herpetologica! Society Page 117 Volume 29 Number 3 September 1993 Page 118 Bulletin of the Maryland Herpetological Society Right groin of S. teapertsis, UCM 28796, male, 59 mm s-v, Cqyame, Lake Catemaco, Veracruz. Thigh to the right. Volume 29 Number 3 September 1993 uiffl o gs 8 « § ° "S ^ 0 « ° § fe C «G *V co ^ o *>gjB| ° 4) O 3 Cco es ^ « b $^5~& i!^i.§2* | o « 6 *? « as SitiHl*. e S «r n 8 c „ » D“u2a5Sa s> fl) -»-> Vj C f-T . —2 i t! p j £ i» ^ “13 -2- c“-a g ^ «0 o g-g* U CO T3 CO a j? ^ £iU S§3 Kl^”l"g ISge-S^l! * & a S E 5 G O *3 a 8 M) S g 't» _ C TJ ^ <» o « 73 - o g » §73 % e go I « 51 .C 3 £ £ ° o • u « w 0^0 Ess •B5 BP <5 73 JB 5 u "O co > DjvhJ M l>rnniv IVHT tus; the role of venoms in food acquisition and defense; and the evolution of venomous squamates. The bode also includes a retrospective essay by Daniel D. Beck, author of the most recent studies on the ecology and behavior of Heloderma , which evaluates the Bogert and Martin del Campo monograph. Dale Belcher has compiled a comprehensive index The color frontispiece is copied from a watercolor of a Gila monster drawn by the noted wildlife artist, David M. Dennis. A special edition of this print, suitable for framing (measuring 9 by 1 1 inches) and limitedlo 100 numbered copies, isavailable; these have been individually signed by Charles Bogert and David Dennis Specifications; 282 pages, 4 portraits of authors, 58 photo¬ graphs. 35 figures, 5 tables, 2 maps. Clothbound in library-grade buckram, format 7 by 10 inches. To be published August 1993. To Order Send orders to the SSAR Publications Secretary, Robert D. Ald¬ ridge, Department of Biology, St. Louis University, St. Louis, Missouri 63103, USA ( telephone 3 14-658-3900 or -3916;/nx 314- 658-3117). Make checks payable to “SSAR”; receipt on request only. Overseas customers must make payment in USA funds. Orders may be charged to MasterCard or Visa (account number and expiration date required); a 5.25% bank charge will be added Bogert and MartIn del Campo: Gila Monster and Allies • Price to SSAR members before August 1, 1993 . US$30 • Price to Institutions and Non-members . $38 ( Postage : Add $2 in USA, add $4 for other countries.) Gila Monster Print, signed and numbered, postpaid . S25 Bulletin of the Maryland Herpetological Society Society Publication Back issues of the Bulletin of the Maryland Herpetological Society, where available, may be obtained by writing the Executive Editor. A list of available issues will be sent upon request. Individual numbers in stock are $2.00 each, unless otherwise noted. The Society also publishes a Newsletter on a somewhat irregular basis. These are distributed to the membership free of charge. Also published are Maryland Herpetofauna Leaflets and these are available at $. 25/page. 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