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Commentaries and Reviews COMMENTARIES AND REVIEWS Editorial Comment. - This section has been established as a forum for the exchange of ideas; opinions, position statements, policy recommendations, and other reviews regarding turtle-related matters . Commentaries and points of view represent the personal opinions of the authors, and are peer-reviewed only to the extent necessary to help authors avoid clear errors or obvious misrepresentations or to improve the clarity of their submission, while allowing them the freedom to express opinions or conclusions that may be at significant variance with those of other authorities. We hope that controversial opinions expressed in this section will be counterbalanced by responsible replies from other specialists, and we encourage a productive dialogue in print between the interested parties . Shorter position statements, policy recommendations, book review s, obituaries, and other reports are reviewed only by the editorial staff. The editors reserve the right to reject any submissions that do not meet clear standards of scientific professionalism. Chelonia n Conserv ation and Biology, 2001, 4(1):21 1- 2 16 © 2001 by Chelonian ResearchFou ndation C. percrassa Cope, 1899, a questionable species from Pleis­ tocene deposits of Port Kennedy Cave, Montgomery County, An Overview of the North American Pennsylvania; and C. owyheensis Brattstrom and Sturn, Turtle Genus Clemmys Ritgen, 1828 1959, from Pliocene (Hemphillian) deposits of Dry Creek, a tributary of Crooked Creek, Malhuer County, Oregon, and Upper Pliocene deposits in Twin Falls County, Idaho (Zug, 1969). Several unanswered questions exist about these fossil 1Department of Biology, George Mason University, species. Are they truly members of the genus Clemmys? If Fairfax, Virginia 22030 USA so, what are the relationships between the various fossil [Fax: 703-993-1046; E-mail: [email protected]] species, and between them and the four living species? These are interesting questions that hopefully someday soon The genus Clemmys Ritgen, 1828, encompasses a group will be answered. of four small to medium-sized, semiaquatic, extant turtles Of the living species, C. guttata is reported from the (Fig. 1) confined to the United States, Canada, and Baja Pleistocene (Rancholabrean) of Dorchester County, South California, Mexico: the spotted turtle, C. guttata; wood Carolina (Bentley and Knight, 1993); C. insculpta from turtle, C. insculpta; bog turtle, C. muhlenbergii; and Pacific Pleistocene (lrvingtonian) deposits in Pennsylvania (Hay, pond turtle, C. marmorata (with two poorly defined subspe­ 1923) and Pleistocene (Rancholabrean) sites in northwest­ cies; Seeliger, 1945; Holland, 1994; Gray, 1995; Janzen et ern Georgia (Holman, 1967), Pennsylvania (Richmond, al., 1997). Below, I present a general overview of the genus, 1964), and Tennessee (Parmalee and Klippel, 1981). Early pointing out various questions or controversies that need to Pleistocene (Blancan) skeletal material assigned to C. be researched, and provide a prognosis for survival of the muhlenbergii has been found in western Maryland (Holman, four species. 1977), and fossils of C. marmorata date from the Pliocene Distinguishing Characteristics. - Usually considered (Blancan) of California and Oregon and the Pleistocene one of the oldest genera in the family Emydidae, the genus (lrvingtonian, Rancholabrean) of California (Brattstrom, Clemmys is best characterized by its hingeless, weakly 1953, 1955; Brattstrom and Stum, 1959; Miller, 1971; buttressed plastron; keeled or smooth carapace; short neck; Gustafason, 1978; Dundas et al., 1996). This is not surpris­ upper jaw lacking a prominent notch or cusps; smooth, ing, because the modern emydid fauna of North America in narrow crushing surfaces on the maxilla which lack involve­ general dates from the Pliocene/Pleistocene. ment from the palatine or pterygoid; no contact between the Intergeneric Relationships. -The four living species pterygoid and basioccipital; contact between the angular of Clemmys are members of the family Emydidae (sensu bone and Meckel' s cartilage; the ju gal tapered ventrally, but Gaffney and Meylan, 1988) and, along with the North not contacting the pterygoid; the frontal bone contributing to American species Emydoidea blandingii and the four spe­ the orbit; and webbed toes. Other characters are discussed by cies of Terrapene plus the Old World species Emys orbicu­ Bramble (1974), Bury and Ernst (1977), Ernst and Barbour laris, form the "Clemmys complex" (all other members of (1989), and Ernst et al. (1994). the family Emydidae belong to the "Chrysemys complex"; Fossil Record. - The oldest fossils possibly assignable Ernst and Barbour, 1989; Ernst et al., 1994). This grouping to Clemmys are those of the questionable Cretaceous (the "Clemmys complex") was first proposed by McDowell (Maestrictian) species C. backmani Russell , 1934, from (1964) under the name "Emys complex" based on skull, jaw, Pretty Butte, Slope County, North Dakota (Quammen, 1992), vertebrae, and shell osteology, and has been supported by the and also known from the Paleocene Ravenscrag formation studiesofMi lstead(l969),Bramble(1974),Bickham(1975, of Big Muddy Valley, Saskatchewan (Russell, 1934). Other 1976), Merkle (1975), Bickham and Baker (1979), Gaffney fossils assignable to Clemmys are: C. morrisiae Hay, 1908, and Meylan (1988), Seidel and Adkins (1989), Bickham et from the Bridger Eocene of Grizzly Buttes in southwestern al. (1996), and Burke et al. (1996). Wyoming; C. saxea Hay, 1903, from the Upper Miocene The relationship of the four genera within the Clemmys Mascall beds on Beaver Creek near Crooked River, Oregon; complex has been debated. The plastron of Clemmys is 212 CHELONIAN CONSERVATION AND BIOLOGY, Volume 4, Number I - 2001 Figure 1. Upper left: Spotted turtle, Clemmys guttata (photo by CHE). Upper right: Wood turtle, Clemmys insculpta (photo by James H. Harding) . Lower left: Pacific pond turtle, Clemmys marmorata (photo by CHE) . Lower right: Bog turtle, Clemmys muhlenbergii (photo by Roger W. Barbour) . hingeless and almost rigid (some movement, correlated with absence of a dorsal keel, presence or absence of serrations on the feeble plastron buttresses, is possible at the bridge), that the posterior rim, shape and dimensions of the neural bones, of the other three genera hinged and movable . Clemmys is and shape and dimensions of various scutes) indicate that C. thought to be ancestral to the hinged genera of which Emys marmorata is more similar to C. guttata and C. insculpta is considered primitive, Emydoidea intermediate, and than to C. muhlenbergii, and that C. insculpta and C. Terrapene most derived (Bramble, 1974). All four genera muhlenbergii are very similar. This arrangement is sup­ share an isoelectric focused myoglobin band not found in the ported by a study of mitochondrial DNA variation by Amato Chrysemys complex. On the basis of this distinct myoglobin et al. (1997). However , analysis of the plastral scute formu­ pattern , Seidel and Adkins (1989) suggested dividing the lae suggests C. muhlenbergii forms a separate group from Emydidae into two subfamilies, of which the Clemmys the other three species, and that C. insculpta is most similar complex would form the subfamily Emydinae, and all other to C. marmorata (Lovich et al., 1991). living emydid species would be assigned to the subfamily The two most discordant studies are those of mitochon­ Deirochelyinae , an arrangement first suggested by Gaffney drial ribosomal RNA genes by Bickham et al. (1996), and the and Meylan (1988) on morphological grounds. This parti­ research by Burke et al. (1996) which included behavioral, tion has been further supported by mitochondrial ribosomal life history, morphological, and ribosomal DNA data. Both RNA gene studies of Bickham et al. (1996). sets of authors considered the genus Clemmys to be lntrageneric Relationships. - Relationships within the paraphyletic. genus Clemmys are confusing and controversial. Bickham Bickham et al. (1996) found the genus to be composed (1975) found no apparent intrageneric karyotypic differ­ of three separate clades, necessitating a new generic ar­ ences, but Merkle (1975), in an electrophoretic study of 17 rangement. They found C. insculpta and C. muhlenbergii different protein bands (apparently using the same speci­ closely related and sharing a common ancestor. These two mens) considered C. guttata and C. muhlenbergii most species formed a sister clade to one containing C. marmorata, similar, with C. muhlenbergii next most similar to C. Emydoidea blandingii and Emys orbicularis. Clemmys marmorata, and C. insculpta the most primitive member, guttata, however, was yet another monospecific clade. Their ·sharing the least numberof protein bands with the other three data supported name changes within the present genus species. This arrangement is supported by the case of hybrid­ Clemmys that would result in the generic name being only ization between wild C. guttata and C. muhlenbergii re­ conserved for C. guttata, the type species of the genus. The ported by Ernst (1983). However, some aspects of my species C. insculpta and C. muhlenbergii would be assigned observations of carapace morphology (shape, presence or to either the genus Calemys Agassiz, 1857, or Glyptemys COMMENTARIES AND REVIEWS 213 Agassiz, 1857, depending on the first revisor, and C. Normally, the spotted turtle is only active from Febru­ marmorata and Emydoidea blandingii would be synony­ ary or March (depending on latitude) to June, with some mized
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