250.1

REPTILIA: TESTUDINES: RHINOCLEMMYS ANNULATA

Catalogue of American Amphibians and . coloration is extremely variable, ranging from unicolored black to dark brown with orange pleural and vertebral blotches to tan ERNST, CARLH. 1980. Rhinoclemmys annulata. with yellow pleural and vertebral blotches. The pleural blotches are often radiations from the dorsa-posterior corner; the vertebral (Gray) keel is usually yellow. The unhinged plastron is well developed, Rhinoclemmys annulata upturned anteriorly, and notched posteriorly; it is a black or dark Brown land terrapin brown with a yellow border and, in some, a yellow midseam. The head is small and has a slightly projecting snout with a slightly Geoclemmys annulata Gray, 1860:231. Type-locality, "Esmeral• beaked upper jaw. A yellowish or reddish stripe extends from the das, Ecuador." Syntypes, juvenile, British Mus. (Natur. orbit at a slight angle to the nape. A stripe runs from the lower Hist.) 1946.1.22.56; adult male, BMNH 1947.3.5.58 collected posterior corner of the orbit to the tympanum where it joins by M. Frezer, no dates; and adult female, BMNH another stripe from the upper jaw. Another stripe passes from 1947.3.5.59, collected by Rosenberg, no date (examined J. the upper anterior corner of the orbit to the tip of the snout. The by author). chin may be mottled with very small dark spots. The feet are Rhinoclemys annulata: Gray, 1863:183. unwebbed or only slightly so. The forelegs have large scales and Chelopus annulatus: Cope, 1865:186. are yellowish with dark stripes of wide black spots. The skin Clemmys annulata: Strauch, 1865:82. elsewhere is gray-brown to yellow. Males have concave plastrons Rhinoclemmys annulata: Gray, 1873:145. First use of combina• tion. and longer tails with the vent beyond the carapace margin; fe• males have flat plastrons and shorter tails with the vent beneath Chelopus gabbii Cope, 1875:153. Type-locality, (Sipurio) Costa the carapace. Rica (from Nat. Mus. Natur. Hist. catalog). Holotype, adult male, U.S. Nat. Mus. 45905, collected by W. N. Gabb, no • DESCRIPTIONS. General descriptions of adults are given by date (examined by author). Boulenger (1889), Siebenrock (1909), Medem (1956, 1962), Prit• gabbi: Gunther, 1885:7. chard (1967), and Ernst (1978). Parsons (1960, 1968) describes the Nicoria gabbii: Boulenger, 1889:119. choanal structure, and the rostral pores and mental glands are Nicoria annulata: Boulenger, 1889:120. described by Winokur and Legler (1974, 1975). annulata: Siebenrock, 1909:499. Geoemyda gabbii: Siebenrock, 1909:499. • ILLUSTRATIONS.Medem (1962) presents a color photograph Rhinoclemmys gabbii: Allee, 1926:451. of an adult; other photographs and drawings of adults are in Med• Geoamyda annulata: Park, Barden, and Williams, 1940:122. Lap• em (1956, 1962), Wermuth and Mertens (1961), Mittermeier sus calami. (1971). Winokur and Legler (1975) show a photograph of a section Geoemyda annulaia: Medem, 1958:19. Lapsus calami. through a mental gland. Carapacial damage by ticks is illustrated Callopsis annulata: Smith, Smith, and Sawin, 1976:216. by Schmidt (1946). • CONTENT. Rhinoclemmys annulata is a monotypic . • DISTRIBUTION.Rhinoclemmys annulata occurs from south• eastern Honduras (Los Andes, Colon) southward through eastern • DEFINITION. Adults grow to 204 mm in carapace length, no Nicaragua, Costa Rica and Panama to western Colombia and size dimorphism. The high carapace is roughened due to growth Ecuador. rings on scutes, posteriorly serrated, and usually widest and high• est just behind the middle. The vertebral scutes are often flat• • FOSSILRECORD. None. tened, but still have a low central blunt keel. The carapacial • PERTINENTLITERATURE. The best general accounts are given by Medem (1956, 1962) and Mittermeier (1971). Other im• portant references are listed by topic. : McDowell -.. (1964), Ernst (1978). Parasites: Fairchild (1943), Schmidt (1946), o 300 400 Evans (1947), Fairchild et al. (1966), Mittermeier (1971), Ernst L '------' ~o MI. , and Ernst (1977). Daily activity pattern: Park et al. (1940). Cer• 600 800 KM. vical vertebrae: Williams (1950). Blood: Frair (1977). Reproduc• tion: Ewert (1979), Moll (1979). '.i • ETYMOLOGY.The name annulata is derived from the Latin annulatus meaning furnished or ornamented with a ring and probably refers to the growth rings on the carapacial scutes. COMMENT Rhinoclemmys annulata is highly variable but not composed of separate recognizable populations. Much variation occurs in all populations regarding carapacial coloration and patterns. The situation is similar to that within populations of the box , Terrapene c. carolina. The species needs to be studied through• out its entire range. Despite being common in some forested areas of Panama, this species has been little studied in the field.

LITERATURECITED Allee, W. C. 1926. Distribution of in a tropical rain• forest with relation to environmental factors. Ecology 7(4):445-468. Boulenger, George Albert. 1889. Catalogue of the chelonians, rhynchocephalians, and crocodiles in the British Museum (Natural History). London. x + 311 p. Cope, Edward D. 1865. Third contribution to the herpetology of tropical America. Proc. Acad. Natur. Sci. Philadelphia / 17:185-196. .i - ••_ ••J ••••• ,/ 1875. On the Batrachia and Reptilia of Costa Rica. 1. Acad. ./ ,/ Natur. Sci. Philadelphia 8(2):93-154 . / Ernst, Carl H. 19'18. A revision of the Neotropical turtle Callopsis (Testudines: Emydidae: Batagurinae). Herpetolog• MAp. The solid circle marks the type-locality; open circles in• ica 34(2): 113-134. dicate other selected localities. -, and Evelyn M. Ernst. 1977. Ectoparasites associated with 250.2

Neotropical of the genus Callopsis (Testudines, Moll, Edward O. 1979. Reproductive cycles and adaptations, Emydidae, Batagurinae). Biotropica 9(2):139-142. p. 305-331. In M. Harless and H. Morlock (eds.), Turtles. Evans, Howard E. 1947. Notes on Panamanian reptiles and Perspectives and research. Wiley-Interscience, New York. amphibians. Copeia 1947(3):166-170. Park, Orlando, Albert Barden, and Eliot Williams. 1940.. Stud• Ewert, Michael A. 1979. The embryo and its egg: development ies in nocturnal ecology, IX. Further analysis of activity of and natural history, p. 333-413. In M. Harless and H. Mor• Panama rain forest animals. Ecology 21(2):122-134. lock (eds.), Turtles. Perspectives and research. Wiley-In• Parsons, Thomas S. 1960. The structure of the choanae of the terscience, New York. (Testudines, Testudinidae). Bull. Mus. Compo Fairchild, G. B. 1943. An annotated list of the bloodsucking Zoo!. 123(4):113-127. insects, ticks and mites known from Panama. Amer. J. Trop. - 1968. Variation in the choanal structure of Recent turtles. Med. 23(6):569-591. Canadian J. Zoo!. 46(6):1235-1263. -, Glen M. Kohls, and Vernon J. Tipton. 1966. The ticks of Pritchard, Peter C. H. 1967. Living turtles of the world. T. F. Panama (Acarina: Ixodoidea), p. 167-219. In R. L. Wenzel H. Pub!., Inc., Jersey City, New Jersey. 288 p. and V. J. Tipton (eds.), Ectoparasites of Panama. Field Mus. Schmidt, Karl P. 1946. Turtles collected by the Smithsonian Natur. Hist., Chicago, Illinois. Biological Survey of the Panama Canal Zone. Smithsonian Frair, Wayne. 1977. Turtle red blood cell packed volumes, Misc. Coll. 106(8):1-9. sizes, and numbers. Herpetologica 33(2):167-190. Siebenrock, Friedrich. 1909. Synopsis der rezenten SchildkrO• Gray, John Edward. 1860. Description of a new species of Geo• ten, mit Beriicksichtigung der in Historischer zeit Ausger• clemmys from Ecuador. Proc. Zoo!. Soc. London 1860:231• storbener Arten. Zoo!. Jahrb., Supp!. (10, pt. 3):427-618. 232. Smith, Hobart M., Rozella B. Smith, and H. Lewis Sawin. 1863. Notes. on American Emydidae, and Professor Agas• 1976. The generic name of the Neotropical semiterrestrial siz's observations on my catalogue of them. Ann. Mag. Nat• emydine turtles. Trans. Kansas Acad. Sci. (1974) 77(4):211• ur. Hist. ser. 3, 12:176-183. 217. 1873. Notes on . Ibid. ser. 4, 11:143-149. Strauch, Alexander. 1865. Die Vertheilung der Schildkroten Giinther, Albert C. L. G. 1885-1902 (1885). Reptilia and Ba• iiber den Erdball. Ein zoogeographischer Versuch. Mem. trachia, p. xx + 326. In F. D. Godman and O. Salvin, Biolo• Acad. Imp. Sci. St.-Petersbourg ser. 7, 8(13):1-207. gia Centrali-Americana. Dulau and Co., London. Wermuth, Heinz, and R. Mertens. 1961. Schildkroten. Kro• McDowell, Samuel B. 1964. Partition of the genus Clemmys kodile. Briickeneschsen. G. Fischer, Jena. 422 p. and related problems in the taxonomy of the aquatic Testu• Williams, Ernest E. 1950. Variation and selection in the cer• dinidae. Proc. Zoo!. Soc. London 143(2):239-279. vical central articulations of living turtles. Bull. Amer. Mus. Medem, Fred. 1956. Informe sobre reptiles Colombianos (I). Natur. Hist. 94(9):507-561. Noticia sobre el primer hallazgo de la tortuga Geoemyda an• Winokur, Robert M., and John M. Legler. 1974. Rostral pores nulata (Gray) en Colombia. Caldasia 7(34):317-325. in turtles. J. Morpho!. 143(1):107-120. 1958. Informe sobre reptiles Colombianos (II). El conoci• -, and - 1975. Chelonian mental glands. Ibid. 147(3):275• miento actual sobre la distribucion geographica de las Tes• 292. tudinata en Colombia. Bo!' Mus. Cienc. Nat., Caracas 2/3 (1-4): 13-45. C. H. ERNST, GEORGEMASONUNIVERSITY,FAIRFAX,VIRGINIA 22030. 1962. La distribucion geographic a y ecologia de los Cro• codylia y Testudinata en el Departamento del Choco. Rev. Primary editor for this account, George Zug. Acad. Colombiana Cienc. Exactos Fis. Natur., Bogota 11(44):279-303. Published 13 August 1980 and Copyright 1980 by the SOCIETY Mittermeier, Russell A. 1971. Notes on the behavior and ecol• FORTHE STUDYOF AMPHIBIANSANDREPTILES. ogy of Rhinoclemys annulata Gray. Herpetologica 27(4):485• 488.