RE- TESTUDINE& EMYDIDAE TERRAPENE CAROLINA Catalogue of American Amphibians and ReptI1es. Tarrudo uirgulataLatreille,In Sonnini and Latreille, 1801:lOO. Type- locality, 'Charleston [Charleston County], South Carolina'. Hol- Ernst, C.H., adJ.P. McBrm 1991. Termpene cadim otype, undesignated Tarado caroliniana Daudin, 1803:207. Lupsus calami Terrapent?caroHna (Ihmaeus) Emys clausa: Schweigger, 1814315. Eastern Box Turtle Emys uirgulata. Schweigger, 1814:316. Emysschneiden' Schweigger, 1814:317. Type4ocaliry, uhwn,re- Tarado carolina Linnaeus, 1758:198. Type-locality, 'Carolina", re- stricted to the vicinity of Philadelphia, Philadelphia County, stricted to vicinity of Charleston, Charleston County, South Car- Pennsylvania by Schmidt, 1953:94. Holotype, undesignated. olina by Schmidt, 1953:93. Holotype, undesignated. Tarupme chusa: Menem, 1820:28. Tarado carimta Linnaeus, 1758:198. Type-locality, 'Calidus regi- Monoclida bmtwkensis Wiesque, 1822:s. Type-locality, 'Ken- onibus", restriaed to Charleston, Charleston County, South Car- tucky". Holotype, undesignated. olina by Schmidt, 1953:93. Holotype, undesignated. Cisado claus(~'Say,1825214. Tarado Mcaudata Lacc5pMe, 1788:169. Nomen substitution pro Tarupme cmlim:Bell, 1825:m. Fituse of combition. ~aradocarolina ~innaeus,1758. Tarupem maculata Bell, 1825:309. Type-locality, unknown, desig- Tarado incamata Bornateme, 178929. Type-bcality, L'America nated as the vicinity of Philadelphia, Philadelphia County, Septentrionale", resuicted to Philadelphia, Philadelphia Coun- Pennsylvania by Schmidt, 1953:s. Holotype, undesignated. ty, Pennsylvania by Schmidt, 1953:93. Holotype, undesignated. Terrupem nebulosa Bell, 1825:310. Type-locality, unknown, desig- Tarado incanerato-striata Bonnaterre, 1789:29. Type-locality, nated as the vicinity of Philadelphia, Philadelphia County, 'L'America Septentrionale", restricted to the vicinity of Philadel- Pennsylvania by Schmidt, 1953:s. Holotype, undesignated. phia, Philadelphia County, Pennsylvania by Schmidt, 195593. Emys hiahhxlata: A. Saint Hilaire, In Cuvier, 1829:ll. Nomen Holotype, undesignated. nudurn Tksado clmrsa Gmelin: 1789:1042. Type-locality, 'Nonh America", Emys(Cisado) carolinas: Gray, 1831a:7. Ex emxe. restricted to the vicinity of Philadelphia, Philadelphia County, Cisado carolina: Gray, 1831b:18. Pennsylvania by Sdunidt, 1953:s. Holotype, undesignated. Emys kimstemoidesGny, 1831b:32. Type-locality, unknown, desig-
Map. Solid circles mark type-localities; hollow circles indicate other selected records. Stars indicate selected fossil localities. nated as the vicinity of Philadelphia, Philadelphia County, of brwacontrast than those of females. The posterior lobe of the Pennsylvania by Schmidt, 1953:94. Holotype, undesignated; plastron is nodyconave in males, flat a slightly convex in Boulenger (1889:117) stated that it was in the collection of the females. Malehindclawsue stocky, considerablycurvedandusually Royal College of Surgeons, London. longer thvl those of fermles, which have shorter, more slender and Cis& viqinsa Agassiz, 1857445. Type-locality, 'New England, straighter hind claws. Males have longer, thicker tails with the vent and westw ard...Michigan ... Cardinw', restricted to Sprinef~eld, near or beyond the posterior carapacial rim; the fdetail is short Middlesex County, Massachusats by Smith and Smith, 1980: with the vent beneaththe posterior marginals. The often usedredeye 553. Syntypes, Museum of Comparative Zoology, 1526,1531- coloration to distinguishthe sexes is not suffiient, as only about 40- 1533 (males), 1527-1530 (fdes), Goldsborn, Wayne Coun- 50%of the males of the subspecies occurring inthe United Stateshave ty, NonhCuoli, colleaed by W. C. Kerr, dateunknown; 15%- red uiOoseph P. Ward, in Litt.). 1535(males), Beaufort County, North Carolina, collected by R. Barnwell, Jr., date unknown; 1537 (male), Ann Arbor, Wash- ~ptions.Gend descriptions arc in Bouleng- (1889, tenaw County, Michigan, collected by Prof. A. Winchell, date 1895), Cahn (1937), Pope (19391, Cur (1952), Mastad (19691, unknown; 1538 (male), Hidunan's Landing, Fulton County, Nietzke (1%9), Cochrvl and Goin (1970), Ernst and Barbour (1972, Kentucky, collected by A. Hyatt, Jr., date unknown; 1539 (fe- 1989). Convlt (1975), Zappalorti (1976), Behl- and King (1979). male), NewJersey, colleued by A. Mayor, dateunknown; 1540 Pritchard (1979), Smith and Smith (19801, Smith and Brodie (19821, (female), New York, collected by J. Froebel, date unknown; Miiller (1987), Clpula (19891, Dundee and Rossman (19891, 1542 (female), Massachusetts, collector and date unknown, (1990), and Comt and Collins (1991). Other descriptions are as 133-1544 (females), collected by Mr. Whittaker, date un- follows: egg (Agassiz, 1857; Cur, 1952; Ernst 4&bur, 1972)) known; 1545,1547-1549(females), 1546,15H)(males), Spring- embryological development (EwM, 1985); hatchlings (Cm, 1952; field, Middlesex County, Massachusetts,collected by J. A. Allen, Ernst and Bubour, 1972); nests (Allad, 1948); kvyotype (St&, dateunknown; 1552 (young male), NorthCarolina, collected by 1972; Webrew, 1977'); courtship (Carpenter and Ferguson, 19n W.C. Kerr, dateunknown; 1554 (juvenile), Delaware River, col- ~mt,1981); skull (Baur, 1891; Ruckes, 1937; McDowell, 1964; lector d date unknown; 1555(1) (juvenilefemale), 1555 (2) GIffney, 1979); cervical venebne mi,1950); pelves (Ruckes, (juvenile male), Washington, D.C., colleaed by Prof. S. F. Baird, 1929); and penis (Zug, 1966). date unknown; 15% (two juveniles, possibly females), locality, collector and date unknown; and 1557 (juvenile, possibly fe- Illustrations. Color illustrations of juveniles and/or adults male), Tennessee, collector d date unknown (series not ex- are in Cochran and Coin (1970), Ernst and Barbour (1972). Conant amined by authors). See Comments. (Ips), Zappalorti (1976), Behler and King (19791, ~ritchd(1979), Tmapne carinakz Strauch, 1862:%. Smith and Brodie (1982), Mirller (1987). Capula (19891, ~undeeand Tmapneeu~iaCop,1869:124. Type-locality, 'Pleistocene de- RW- (19891, and co-t and co~i(1991); bkck and whjte posits in [Oxford Neck] Talbot County, Maryland". Holotype, photographs or drawings are in Sowerby and Leu (18721, Cahn American Museum of Natural History 1484, entoplastron and (1933, Pope (19391, Car (19521, Wermuth and Mertens (1%1), posterior carapace element (not examined by authors). Milstead (1%9), Nietzke (1%9), Ernst and Bubour (1972, 1989)~ Tmapnepulnami Hay, 1906:30. Type-ldty, 'Alifia River, Flori- Zappdorti (1976), Pritchard (1979), Smith and Smith (19801, Obst da, about a mile from its mouth". Holotype, American Museum (1986), d Tyning (1990). Other illustrations are as follows: eggs of Natunl History 6097, left hypoplastrd bone, collected by and embryos (Agassiz, 1857; Ewert, 19791, skull (Boulenger, 1889; Professor F.W. Putrum (nu examined by authors). Auffenberg, 1958; Wennuth and Mertens, 1961; Emst and Bubour, Tarapnefonnara Hay, 1916a:57. Type-locality, 'Ocala, Marion 1972;~Iffney, 19791, pelves (Ruckes, 19291, vestigial ribs (Hoffsten- County], Florida. Pleistocene". Holotype, United StatesNational and Gax, 1%9), brain (Starck, 1979), ear and its functions (Wever, Museum 8825 (formerly Geological Survey of Florida 2973), 1978), oxygen affmity and discciation curves (Dessauer, 1970; posterior two-thirds of anpace, collector and date not given Wood and Lenfant, 1976), sexual dimorphism (Cahn, 1937), court- (examined by authors). ship and mating behavior (Cahn, 1937; Carpenter and Ferguson, Tmapmcardli~' Proctor, 392220. E* enuw. 1977; Obst, 1986 andMuller, 1987), nesting behavior (Akud, 19351, T- cawlina: Breder, 192463. Ex m. plaster cws of nests (Allad, 19481, copra lutes Wtland, 19511, TmupmsingletoniGilmore,1928: 1 .Type-locality, 'Twomileswest resting form (Stickel, 1950), and distribution Overson, 1986). of Melbourne, Brevard County, Flori&... Pleistocene." Hole type, U.S. National Museum 11181, partial carapace, presented Distribution. In the United States, Tmupm camlina by C.P. Singleton in either 1925 or 1926 (examined by authors). ranges from southern Maine south to the Florida Keys and west to TmapmecawIinap~mi:Auffenberg,1958:53. Fiuseofrecog- southern Michigan, Illinois, eastern Kansas, Oklahoma, and Texas. nized combination The species may also be found in isolated localities in New York and western Kansas. In Mhico, it occurs in the states of Campeche, Content. Six living subspecies(T~~~~cawIi~cadinqQuintano Roo, Sm Luis Potosi, Tunaulipas, Veracruz, and Yucarh. T. c. mericuna, T. c. hiunguis, T. c. major, T. c. bun, and T. c. The peripheral boundaries of the species range have been somewhat yucalrma), and the fossil subspecies T. c.putmami are recognized. douded because this turtle is a popular pet and individuals are often transported and then released in places far from their natural distri- DefMtIon. Ten;apsnecamlinahas a keeled, domed, elon- butional area. gated carapace (to 21 cm) which is not serrated posteriorly and is highest posterior to the plvtnl hinge. The medial keel usually is Possa Record. Now assigned to TBIT~~~cadi- are the present on vertebrals 24, vertebral 1 is elevated at a steep angle (SO0 extinct subspecies T. c.putnami Hay, 1906 and several fossils once or more), and the fitmargin& usually are reaulgular. Vertebrals considered swcies (Milstead 1%9): Cistudo mamockico~e.1878: are wider than long. The carapace usually is brown but also often horn-colored or black, usually with an exwemely variable pattern of yellow or orange radiating lines, spots, bars, or irregular blotches on ~ay,i916b; T. ~ay,1921; T. i+Aw H;~, 1924; 2 each scute. The plastron bears a moveable hinge between the singktoni Gilmore, 1928; and Tracbemys tEucbocarinata Hay, pectoral and abdominal scutes, and is often as long or longer than the 1916a. carapace. An Mary scute may be present. Usu;tlly, the hindlobe Fossil remains of T. cmlina are known from the Miocene does nuhave a posterior notch. The plastron is tan to brown; it may (Cluendonid of Kansas and Nebraska (Hdmul. 1975.1981). Plie be patternless, show dark blotches or smudges, a dark central area &ne (~ernphiuianand Blanan) of Flor& (Miliead, 1%9),ud in with branches along the seams, or have a pattern of yellow ndiating the Pleistocene from the Blanan of Florida and Muvland (Milstead. lines. The head is small to moderate in size with a nonprojecting 1969; Holman, 19n); Wingtonian of Florida, &, Maryland; snout and a medially hooked upper jaw lacking a notch Its dorsal Nebraska and Texas (Milstead, 1969), d the Rancholabrean of surfaceis covered withsmoothskin. Colorationusuallyis brownwith Florida, Georgia, Mississippi, Missouri, Texas, and Virginia yellow, orange, or red latenl spots, but the head may be patternless (Auffenberg, 1957, 1959; Milstead, 1%9; Holman, 1967, 196%, or entirely reddish, white, brown or black. The forelimbs are covered 19694 1978; Mdure and Milstead, 1967; Gillene, 1974;Jackson and with large brown, yellow, orange or red scales; other skin usually is Kaye, 1974; Moodie and Van Devender, 19nFay, 1984; Holman and brown. The toes are heavily clawed and partially webbed; hind toes Clausen, 1984; Holman and Winkler, 1987). Remains of T. camlina total either three or four per foot. The foreleg colors of most males are have also been found at archedogid sites; summvies of such mords in Canah, the United States, and Mexico ue in Hatt and cvapacial pining (Carpenter, 1956); immunology and electrophore- hngebutel(1953), ~leakney(1958),W~(1%5), Adler(l%8,1%9, sis (Leone d Wilson, 1961; Frair, 1962,1963Hildeman, 1962; Dozy 1970), Lztham (1%9b), and Nvvez (1976). et al., 1964; Cooper et al., 1985; Mao et al., 1987); physiology (Anderson d Wilbur, 1948; Altland and Parker, 1955; Hunon and perttoeat Il- General accounts are in Baur (18931, Goodknight, 1957;Pukerand Altland, 1957; Payne, 1957;Altlandand Taylw (1895), Brimley (1904), Babcock (19191, Dims (19341, Thompson, 1958; Musacchia and Chladek, 1961; Smith and Nickon, AUard (1935, 1948, 19491, Clhn (19371, Pope (1939)) can (1952)a 1961; Schwvtz and Kaph, 1962; Adler and Small, 1963; Dodge and Dickson (1953), Oliver (19551, Duehand Schwartz (1958)s Folk, 1963; Robiion and Schmidt-Nielsen, 1963; Hutton, 1964; Kirsche (1%9), Emst and Barbour (1972, 1989), Conant (1979, Payne and Burke, 1964; Quay and Wioft, 1964; Hutchison and ~urphy(1976), Hatless and Morlock (19791, Pritchard (19791, Smith Szarski, 1965; Schmidt-Nielsen and Bentley, 1966; Helmy and Hack, and smith (1980), Stuan and Miller (1987) and Dundee and k~man 1967; SullivanandRiggs, 1967; Ernst, 1968; Helrny etal., 1969;Morgan (1989). Other papers are listed by topic u follows: evolution and Singh, 1969; Dessauer, 1970; Brisbi, 1972; Mogareidge and (~ilstead,1969; Gillene, 1974); systematics wbour and Stetson, Hamrnel, 1975; Bennett and Dawson, 1976; Bentley, 1976; 1931;Milller, 1936; Smith, 1939; Smithand~aylor,1950b;~uffenberg, Elghammer andJohnson, 1976; McDonald, 1976; Wood and Want, 1958; Mutin, 19%; McDowell, 1964; McCoy and Richmond, 1966; 1976; Skoctyiu, 1978; Belekhova, 1979; Enkine and Hutchison, Milstead, 1969; Smith and Smith, 1980; Kiester et al., 1982); karyotype 1981; Sturbaum, 1981; Sturbaum and Berg- 1981; Bartholomew, (Huang and Clark, 1967; Clark et al.,1970; Stock, 1972; Stock and 1982; Bennett, 1982; Gregory, 1982; Lillywhite andMaderson, 1982; Mengden, 1975; Killebrew, 1977; Bickhvnand Baker, 1979; Bidham Minnich, 1982; Srurbaum, 1982; Monagas and Ganen, 1983; Karasov and Carr, 1983); distribution and zoogeography (Smith and Taylor, et al., 1985; Risher and Claussen, 1987; Packard and Packard, 1988; 1950a, 1950b; Neill, 19%, 1958; Auffenberg and ~ilstead,1965; Uluchand Anderson, 1988; Vivien-Raels et al., 1988; Olson, 1989;Yi Savage, 1966; Blaney, 1971;~verson, 1986; auks st is and Juuen, 1988; et al., 1989; Costanzo and Chussen, 1990; Claussen et d., 1991); Iverson and Etchber~er,1989; Pdley, 1989; Gibbons and Semliuch, parasites and disease (Kepner, 1912; Ewing, 1926; Peters, 1948; 1991); morphology 6oohnston, 1915;~uckes, 1929; Knoll, 1935; Lynn, Mehrtens, 1951; King and Griffo, 1958; Jackson et al., 1969, 1972; 1937; Haines, 1946, 1969; Green, 1950; Altland, 1951; Hebard and Emst and Barbour, 1972; Pence and Casto, 1975;Abercrornbie, 1977; Chariiper, 1955; Zvlgerl and Johnson, 1957; Smith, 1958; Smith and Emst andEmst, 1977,1979; Lawler, 1977; CrispensandMarion, 1981; J-=, 19%; pusom, 1960, 1%8; Richmond, 1964;Wii-andHan, Rowley andMoskduk, 1985; Wejumvlnet al., 1985;McAllister, 1987; 1964; Sehe, 1965; Zug, 1966, 1971; Hofstetter and Gasc, 1969; Baxter and Meek, 1988; Oliver et al., 1988); DDT and other con- Milstead, 1969; Zangerl, 1969; Baird, 1970; Bockman, 1970; Carey, taminants (Stickel, 1951; Ferguson, 1963; Holcomb and Parker, 1979; 1970; Duguy, 1970; Saint-Girons, 1970; Cook et al., 1972; Beresford et al., 1981); radiation effects (Altland et al., 1951; Schurmcher, 1973; Walker, 1973, 1979; Winokur and Legler, 1974; Cmgrove, 1965); enrmpment by railroad tracks (Helms and Stains, FOX,1977; Parsons and Cameron, 1977; Wever, 1978; Elgh-r et 1966); ecology (Surface, 1908; Weunore, 1920; Nichols, 193%; Penn d., 1979; Gaffney, 1979; Stuck, 1979; Buten, 1981; Leuck and and Potthust, 1940; Bogen and Cowles, 1947; Allad, 1948, 1949; Carpenter, 1981; Wibern and Ingold, 1983; Holman, 1984; Wiokur, Neill, 1948b, 1958; Barbour, 1950; Stickel, 1950,1951; Myers, 1952; 1988); embryology and development (Nelson, 1953; Crooks and Wood and Goodwin, 1954; Guibe and Saint-Giro-, 1955; Deuoff, Smith, 1958; Tucker and Funk, 1976; Ewen, 1979,1985; Bellairs and 1956; ~utchison,1956; Cupenter, 1957; Williams, 1962; Murphy, Bryant, 1985; Raynaud, 1985; Cohen, 1986; Rose, 1986); histology 1964; Branstrom, 1965; Hutchison et al., 1966; ~dler,1968; Emst. (Balmer, 1951; Clark and Karzon, 1%7a, 1967b; Huang and Chk, 1968; Dolbeer, 1969,1971; McCoy, 1969; Russo, 1973; Reagan, 1974; 1967; Parsons and Cameron, 19m, growth (L~M,1937; Babcock, Schwartz and Schwutz, 1974; Yahner, 1974; Dodge et al., 1978; 1939; Ewing, 1939; Nichols, 1939a; Panerson, 1978; Stickel and Stickel, 1978; Auffenberg and Iverson, 1979; Hutchison, 1979; Cur P Bunck, 1989); longevity flowmend, 1926; Deck, 1927; Babcd, andHouseal,1981; EnkineandHutchison, 1981;RustandRoth, 1981; 1928;~llard,1949; Edney and Allen, 1951; Price, 1951; Lapeer, 1953; Avery, 1982; Clark, 1982; Gregory, 1982; Iverson, 1982; Stnss et al., Oliver, 1953a, 1953b; Nigrelli, 1954; Junes, 1961; Lthm and 1982; Wibern and Ingold, 1983; Strang, 1983; Holman, 1984; Muir, Schlauch, 1969; Graham and Hutchison, 1969; Bowler, 1977; Wi- 1984; Schwvtt et al., 1984; Kiester, 1985; Vance, 1985; Maun and liams and Parker, 1987); albinism (Hensley, 1959; Am&, 1980); Brooks, 1987;Gatten, 1987; Wiand Parker, 1987;Greene, 1988;
Figure 1. United States subspecies of Terrapme cadinq left to right: bauri, camlina, hiunguis, and mapr(Roger W. Bubour).
k 1951; Evans, 1951a. 1951b, 1952,1953; Legler, 1955; Smith and List, 1955; Fox, 19%; Lin, 1958; Ev-, 1961, 1%8; Schwm, ~aens,1973; Hamand Gist, 1975; Tucker and Punk, 1976,1977, 1978a, 1978b; Carpenter and Perguson, 1977; Iverson, 1977; Congello, 1978; Meld,1979; Ew-, 1979; Moll, 1979; Berry and Shine, 1980; Reimer, 1981; Wuner, 1982; Din41983; agiol la, 1984; Levell, 1985; Congdon and Gibbons, 1985; Ewa, 1985; binsky et d., 1987; Packard and Packad, 1988); hybridization (cluk, 1935; shvlnon and Smdh, 1949; Menens, 19% 1956; Smith, 1955; Blaney, 1968; Ward, 1968); intergradation (Smith and mders, 1952).
Etymology. The specific name cumlina is for the Carolina region of North Arneria. Subspecific names uederived as follows: mg*icrma is for Mhico; triunguis is from the Latin words hi, three, and unguiculus, nail or daw, refering to the three toes on each hind foot; major is Latin for larger or greater, and refers to the relatively large size of this subspecies; bauri is a patronym for George Herman Figure 2. T811apne CU~~Mcadina (Roger W. Barbour). Carl Ludwig Baur, an early -her of the genus; and yucatuma is for the YUU~region of Mhico. ~dvnset d., 1989; Congdon et d., 1989; Stickel, 1989; Grab-, 1990); food habits (Surface, 1908;Cm.w.r, 1940; Moore, 1953; Myers, Comments. In addition to the syntypes listed for Cistudo 1956; Bush, 1959;Klirnsva and Newsome, 1%0; lathun, 1968,1%91; vi'ginea, a series of ten embryos and hatchlings (Museum of Com Boice, 1970;Alsop and Wallace, 1978; Rust and Roth, 1981; Bnun and parative Zoology, 168; not examined by authors) was figured by Brooks, 1987; Anton, 1990); behavior Wi- 1922; Bred=, 1924, Agwiz (18Sn, but whether or not these specimens should dso be 1927; Frothingham, 1936; Nichds, 1939b, 1939~;Allad, 1948,1949; considered syntypes is questionable. Neill, I%&; Stickel, 1950; Beltz, 1954; Wood and Goodwin, 1954; The holotype of the fossil species Tmapene imprara was not Myers, 1956; Wwer and Vernon, 1956; Gould, 1957, 1959; Belu, designated by Hay (1924), but, since the nuw was published in 1958; Verway, 1958; Hutchison and Vinegar, 1963; Evans, 1%4; combination with an illustration, it does meet the ICZN pre-1931 Hayes and Saiff, 1%7; Rosenbslum, 1%8; Lemkau, 1970; Dolbeer, rules, an4 since the nominal species is based on this single specimen, 1971; Flanigan et d., 1974; Schwvtz and Schwutz, 1974; Brosgole, it is the holotype by default. The type-specimen is part of a collection 1976; Burghudt, 1977; Carpenter and Ferguson, 1977; Shdet, 19n made by Dr. Francis in the early 1900's originally deposited in the Hardiig, 1978; Wever, 1978; bless, 1979; Manton, 1979; Morlock, museum of Texas A&M University. After that museum dosed, the 1979; Posey, 1979; Tyler, 1979; Walker, 1979; DeRosa and Taylor, Francis collection was transferred to the Texas Memorial Museum, 1980, 1982; Cur and Housed, 1981; Erskine and Hutchison, 1981; University ofTexas,Austin, and the specimen is now in the Vertebrate Kiester et d., 1982; Wdbern, 1982; Schwartz et d., 1984; Mathis and Paleontology Laboratory, Bdcones Research Center, of that institu- Moore, 1988; Abet d., 1989; Hayes, 1989; Stickel, 1989); repro- tion (Melissa C. Wims, in Litt.). duction (Ewing, 1933; Nard, 1939; Nichols, 1940; Penn and Syntypes 15051508 of Tmapenec. majorhave been identified Pottharst, 1940; Ewing, 1943; Firan, 1948; NeU, I%&; Abnd, u T. c. bauri(Jose P. Rosado, in Litt.).
Figure 3. Variation in Tmapnecmolina catdiM (Roger W.Barbour). Milstead (1967,1969) considered Tmcamlina to consist Terrapene carinata: Strauch, 1862:96. See species synonomy. of several subspecies, and thought that T. c. majorand T. c. hiunguis ~erra$meeumia Cope, 1869:124. See saessynonomy. intergraded along the Gulf Coast in southeastern Louisii. How- Cisado cmlina w.cimsIemoides: Boulenger, 1889:117. Nomen r' ever, Ward (1980) regarded T. mapras a distinct species ranging subsIitutkm fromthe Mississippi River to peninsular Florida. Ward did not use the T- camlina cadinaStejneger and Barbour, 1917:115. Fii same characters as Milstead in defining T. majw, and several of his use of combiition statements are contradictory (Dundee and Rossman, 1989:194). Terepene camlina: Breder, 192463. See species synonomy. Dundee and Rossman (1989), after examining additional specimens from St. Charles Parrish, Louisiia, concluded that T. c. mapris best Defhition. The carapace is shon, broad, and brightly regarded as a subspecies, as interpreted by Milstead (1969). patterned with the rmrginals nearly vertical or slightly flared, and each hind foot has four toes.
2. Terrapene caroHna mexicana (Gray) Tesado camlina Linnaeus, 1758:198. See species synonomy. Tesado cannala linnaeus, 1758:198. See species synonomy. Cisado(0nycbohfu) marlcanaGny, 1849:17. Type-locality,'Mexi- TesW brsvizmdatah-e, 1788:169. See species synonomy. co', restricted to vicinity of Tunpico, Tamaulipas by MWer, Teshdo incanerala Bonnaterre, 1789:29. See species synonomy. 1936:112. Syntypes, British Museum of Natural History Teshdo incaner(11o-shialaBonnaterre,1789:29. Seespecies synon- 1947.3.5.48, stuffed, mounted male, collector and date un- omy. known; and 1947.3.4.3, stuffed adult female, purchased by 'Mr. Teshrdo clm*sa Gmelin, 1789:1042. See species synonomy. Warwick', date unknown (examined by authors). Teshdo uirguhta htreille, In Sonnini and Latreille, 1801:lOO. See Cistudo maxicana: Gray, 1856:40. species synonomy. Onychohia mgxicana: Dugb, 1888:107. Teshrdo cumliniana Daudin, 1803:207. See species synonomy. Cishdo camlina w. mexicana: Boulenga, 1889:118. Emys clausa: Schweigger, 1814:315. See species synonomy. Onichotria mgxica~Herrera, 1890:330. Lapsus cakami Emys uirgulah Schweigger, 1814:316. See species synonomy. Cbelopus mgxicanus: Vehsco, 1892:79. Emys scbneiden' Schweigger, 1814:317. See species synonomy. Terrapew mexicana. Baur, 1893677. Terrtpm clausa: Merrem, 1820:28. See species synonomy. Terrapewgokimani Stejneger, 1933:119. Type-locality, 'Chijol (or Monoclida kentuckensisRafiisque, 1822:s. See species synonomy. Chijoles), southeastern corner of the State of San Luis Potosi, Cishdo clausa: Say, 1825:214. See species synonomy. Mexico; in the coast plain." Hdotype, U.S. National Museum Terrapew camlina: Bell, 1825:W. See species synonomy. 46251, alcoholic female, collected by E.W. Nelson and EA. Terrapew macuhka Bell, 1825:309. See species synonomy. Goldman, 11 May 1898 (examined by authors). Terrapew nebubsa Bell, 1825:310. See species synonomy. Terrapewyucalana: Difmus, 19344. In part. Emys IritenIaculaIa:A. Saint Hilaire, InCuvier, 1829:ll. See species Terrapew mkoni: Diman, 193440. In pan. synonomy. Terrapew mexicana mcdcunu: Smith and Taylor, 1950b:346. Emys(Cishdo) camlina. Gray, 1831a:7. See species synonomy. Terrapew camlina me-kana: Milstead, 1967: 168. Fituse of com- Cishdo camlina: Gray, 1831b:18. See species synonomy. bination Emys kimstemoides Gray, 1831b:32. See species synonomy. Termmexicatm Nierzke, 1975371. Ex emm?. P Cishdo viqinea Agassiz, 1857:445. See species synonomy. Terapme makana mexic(1~'Nietzke, 1973371. L+DSUS cakami
1 Figure 4. Terrapew cadina mgxicana (Roger W. Barbour).
I i Dellnition. The carapace is elongated and highdomed with the third vertebral elevated into a small hump and only moderate flaring of the posterior marginals; the carapace is pale yellow or tan to light olive with dark seuns; each hind food has three toes.
3. Terrapene cam#na triunguis (Agassiz)
Cisludo hiunguis Agassiz, 1857:445. Type-locality, 'New Orleans', Louisiia. Syntypes, Museum of Comparative Zoology 1519 (seven females, one male), 1522 (female); from type-locality, colleaed by Dr. N.P. Benedia, no date; 15251524 (females), 1525(male) from type-locality, collected by F.C. Copes, no date (not examined by authors); U.S. National Museum86871-86872 (alcoholic females, formerly Museum of Comparative Zoology 1519)fromtype-locality, colleaed by Dr. N.P. Benedia, no date (examined by authors); U.S. National Museum 22 (alcoholic fe- rrrale) from Washington, Adams County, Mississippi, collected by'Wailes",dateunknown(examinedbyauthors); U.S. Nation- al Museum 213736 (alcoholic female) from Missiiippi, collea- ed by 'Wailes", dateunknown(examined by authors); U.S. Nati- onal Museum7546 (dry, stuffed female with separate skull) and 131838 (alcoholic female) localities, collectors, and dates un- 5. Terrapene camtina bauri Taylor known (examined by authors). Cisludo cmlina var. cinoslemoida: Ebdenger, 1889:117. In put. Tmapene baun"raylor, 1895576. Type-loczliy, 'Florid?", re~victed Cis&& mamocki: Cope, 1878:229. In put. to Orlando, Orange County, Florida by Sdunidt, 1953:94. Ho- Terrupm rriunguis. Taylor, 1895: lotype, U.S. National Museum8352, alcoholk female, colleaed Terrupmcatolina hiunguCStrecker, 1910:121. Pituseofcombi- by F.B. Meek in 1875 (examined by authors). nation. T- innaxiaHay, 1916a:61. Type-locality, 'Pleistocene. Vero, Terrapem wbitneyi Hay, 1916b:8. Type-locality, post-Wisconsin St. hcie County, Florida.' Holotype, U.S. National Museum gravels in Austin, Travis County, Texas (from label: 'Basement 8824 (originally Florida Geological Survey 7080), complete car- of Littlefield Buildii, Austin, Texas, Pleistocene"). Holotype, apace, collected by E.H. Sellards, no date (examined by auth- U.S. National Museum 8617, complete carapace and plastron ors). (examined by authors). Teropane camlina bau~Cur, 1940:lOO. First use of combiition. Terrapem bulwdaHay, 192&133. Type-locality, not given. Holo- type, U.S. National Museum 9921, disarticulated posterior por- Deflnltioa. The dark brown carapace has a bright pattern of tion of carapace, and plastron, collected by D.V. Schuchvdt in yellow radiating lines, is highest posterior to the center, and has lntle 1915 (examined by authors). or no king of the posterior margin&. The plastron also beam a n H. buluada: Hav. 1920:1%. Ex enwe. pattern of radiating light lines on each scute, and two light, longinr- ~errapemirnpn&a~a~,;924:245. Type-locality, 'Munson's Shoals, dinal stripes ue present on each side of the head. Each hind food 2 or 3 miles below Pinbridge", Bmzos County, Texas. Hdotype, usually has only three toes. Texas Memorial ~useuGUniversity of Texas (TMM-T~U) 2401, posterior lobe of plastron(apparenlly misidentified as an- terior lobe inoriginaldescription), collectedby Dr. Mark Francis in 1923 (not examined by authors). See Comments. TerrupmlhnmsisOelrich, 1953:35. Type-locality, 'Sangaman, late CfludoyucakmaBoulenger, 1895330. Type-locality, 'Nonh Yuu- Pleistocene ... tributary of Shorts Creek ... Lone Tree Arroyo ... tan', M6xico; restricted to Chichen Itu, Yuath, Mhico by Meade County, Kansas.'' Holotype, Museum of Paleontology, Smith and Taylor, 19501:35. Synrypes, British Museum of Natu- University of Michigan26957, putid cvapaceandplastron, and ral History 1947.3.5.4547,dry adult females, collected by "Gau- other bones, colleaed by University of Michigan Field Pw,28 me?, but origited from 0.Salvin (examined by authors). July 1950 (examined by authors). Cfrudo maicanu: Barbour and Cole, 1906,147. NecGray, 1849. Terrapem canalicuhta: Milstead, 1956:163. In put. Tmapneyucatamz Sibenrock, 1909:492. Tmupmmaicamz Muller, 1936:lOO. NecGray, 1849. Definition. The highest point on the carapace is more Tmapne msricana yucakavkz Smith, 1939:17. posteriorly positioned than in other subspecies of T. camlina; in Tenapene cawlinayucatanm Milstead, 1%7:168. Fituse of cam- uoss-sedion it is peaked in appearance, and is typically tan or olive bition. with an obacure panern of radiating dashes and, often, dark seam Tmqtmw maicanujucatnna: Nietzke, 1973371. Lupus cakami borders. Orange or yellow spots usually are conspicuous on both head and forelimbs;males may have totally red heads. Three toes are Deflnltloa The carapace is long, high-domed, with the third present on each hind foot. vertebral elevated into a small hump, and little flaring of the posterior margin&. It is tan a straw-colorkd with dark radiating dashes or black sateborders. Each hind food has four toes. 4. Terrapene carotina ~jor(Agassiz)
Cistudo majorAgwiz, 1857445. Type-locality, 'Mobile Wbama] ...Florida', restricted to Mobile, Mobile County, Alabama by Aberaombie,J. 1977. Myiasis in the eastern box turtle caused by Schmidt, 1953:94. Syntypes, Museum of Comparative Zoology Pbasniciu c~uiridis(Diptera:Wphoridae). J. Washing- 1505,1507,1509-1510 (females), 1506 (male), 1508 (juvenile, ton Aud. Sci. 67:155-1%. probably male) (not examined by authors). See Comments. Ah,N.A., D.L Clussen, and J. Skillings. 1989. Effeds of tempera- Cisludo cmolina vat. major: Boulenger, 1889:117. ture on vdu~vylocomuion of the -ern box tude, Terra- Terrqtmw major: Taylor, 1895575. pens camlina camlina Copeia 1989:905915. Tmapnecadina major: Cur,194@101. Piuse of combination Adler, K.K. 1968. Tudes from archedogid sites in the Great bkes q region Michigan Archaeol. 14:147-163. DeWtion. The carapace is long, sometimes exceeding 20 -1969. Elimination of box turtles in western New York state by an, with few or no markings, or marks obscured by Mack or tan prehistoric man. J. Herpetol. 2:179. pigment in males, but radiatingyellow spots or dashes in females; the -1970. The influenceof prehistoricmanon the distributionof the posterior marginals are strongly flared, and four toes occur on each box turtle. Ann. Carnegie Mus. Nat. Hit. 41:26f 280. hind foot. -and R.A. Small. 1963. Bioelecuogenesis in the urinary bladders of turtles, genera Aademysand Terrupm J. Ohio Herpad. Baur, G. 1891. American box tortoises. Science 17:19@191. Soc. 4:63-64. -.1893. Furthernotesonhmuicanboxturtles. Amer. Nat. m6n- Agassiz, L 1857. Contributions to the natural history of the United 678. States of America, first monograph. Vol. 1, put 2. North Ameri- mer,J.S. and R Meek. 1988. Cryosurgery in the tmunent of skin can Testudinata. Little, Brown and Co., Boston. p. 233-452d. disorders in reptiles. Herpad. J. 1227-229. Allad, HA. 1935. 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