The Oriental Colubrine Genus Coelognathus Fitzinger, 1843 and Classification of Old and New World Racers and Ratsnakes (Reptilia, Squamata, Colubridae, Colubrinae)

Urs Utiger, Beat Schätti, Notker Helfenberger

Abstract


The phylogenetic relationships of the colubrine genus Coelognathus is examined on the basis of two partially sequenced mitochondrial genes, COI and 12S rRNA. Maximum parsimony reconstructions show that C. erythrurus, C. flavolineatus, C. helena, C. radiatus, and C. subradiatus share a common ancestor. Coelognathus spp. represent an evolutionary lineage among Oriental racers. Based on molecular data, external morphology, and pulmonary features, Coluber prasinus Blyth and Herpetodryas frenatus Gray are referred to Gonyosoma Wagler (four species). Hemerophis from Sokotra (monotypic) is the sister taxon of Palaearctic and Saharo-Sindian racers. Their phylogenetic relationships with Oriental racers are not clear. Old and New World racers belong to genetically independent lineages. The latter includes Salvadora spp. Masticophis Baird is considered to be a junior synonym of Coluber Linnaeus. A new generic name, Oreocryptophis, is proposed for the East Asian ratsnake taxon Coluber porphyraceus Cantor. Coronellini Jan is proposed for Old and New World ratsnakes. The tribe is monophyletic and made up of at least eight Old World (Coronella, Elaphe [sensu stricto], Euprepiophis, Oocatochus, Oreocryptophis, Orthriophis, Rhinechis, and Zamenis) and ten New World (mostly Nearctic) genera (Arizona, Bogertophis, Cemophora, Lampropeltis, Pantherophis, Pituophis, Pseudelaphe, Rhinocheilus, Senticolis, and Stilosoma).

Keywords


Coelognathus; Gonyosoma; Oreocryptophis (nomen novum); Oriental racers; phylogeny; Colubrini; Coronellini; morphology; mtDNA; COI; 12S rDNA

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References


Amaral A. do (1930), «Contribuição ao conhecimento dos ophidios de Brasil. IV. Lista remissiva dos ophidios do Brasil», Mem. Inst. Butant., 4[1929], IV + 69 – 126.

Amaral A. do (1934), «Estudos sôbre ophidios neotrópicos. XXX. Novo gênero e espécie de colubrideo na fauna da Colombia», Mem. Inst. Butant., 8[1933 – 1934], 157 – 159.

Auffenberg W. (1955), «A reconsideration of the racer, Coluber constrictor, in eastern United States», Tul. Stud. Zool., 2(6), 87 – 155.

Baird S. F. (1859), «Reptiles of the boundary, with notes by the naturalists of the survey», in: Emory W. H. (ed.), Report on the United States and Mexican Boundary Survey. Part II. Zoology of the Boundary, Govt. Print Office, Washington, pp. 1 – 35.

Baird S. F. and Girard C. (1853), Catalogue of North American Reptiles in the Museum of the Smithsonian Institution. Part I. Serpents, Smithsonian Inst., Washington.

Bleeker P. (1858), «Gonyosoma jansenii Blkr., eene nieuwe slang van Manado», Nat. Tijdschr. Nederl. Ind., 16, 242.

Blyth E. (1854), «Notices and descriptions of various reptiles, new or little known», J. Asiatic Soc. Bengal, 23(3), 287 – 302.

Blyth E. (1860), «Report of the Curator, Zoological Department», J. Asiatic Soc. Bengal, 29, 87 – 115.

Boettger O. (1886), «Diagnoses reptilium novorum ab ill. viris O. Herz et consule Dr. O. Fr. de Moellendorff in Sina meridionali repertorum», Zool. Anz., 9, 519 – 520.

Boettger O. (1888), «Über die Reptilien und Batrachier Transcaspiens (Vorläufige Mittheilung)», Zool. Anz., 11(279), 259 – 263.

Boie F. (1826), «Generalübersicht der Familien und Gattungen der Ophidier», Isis [Oken], 19(10), 981 – 982.

Boie F. (1827), «Bemerkungen über Merrem’s Versuch eines Systems der Amphibien. 1te Lieferung. Ophidier», Isis [Oken], 20(10), 508 – 566.;

Bonaparte C. L. (1834), Coluber leopardinus. Iconografia della Fauna Italica per le quattro Classi degli Animali vertebrati [1832 – 1841]. II. Amfibi [Fascicolo VII, punt. 38], Salviucci, Roma.

Boulenger G. A. (1893), Catalogue of the Snakes in the British Museum (Natural History), Vol. 1, Trustees Br. Mus. (Nat. Hist.), London.

Boulenger G. A. (1894), Catalogue of the Snakes in the British Museum (Natural History), Vol. 2, Trustees Br. Mus. (Nat. Hist.), London.

Boulenger G. A. (1898), «On a second collection of reptiles made by Mr. E. Lort-Phillips in Somaliland», Ann. Mag. Nat. Hist. Ser. 7, 2, 130 – 133.

Brongersma L. D. (1957), «Notes upon the trachea, the lungs, and the pulmonary artery in snakes. I», Proc. K. Ned. Akad. Vet., 60(C), 299 – 308.

Brown A. E. (1890), «On a new genus of Colubridae from Florida», Proc. Acad. Nat. Sci. Phil., 42, 199 – 200.

Cantor T. (1839), «Spicilegium serpentium indicorum (II)», Proc. Zool. Soc. Lond., 1839(7), 49 – 55.

Clark P. J. and Inger R. F. (1942), «Scale reductions in snakes», Copeia, 1942(3), 163 – 170.

Cope E. D. (1860), «Catalogue of the Colubridae in the Museum of the Academy of Natural Sciences of Philadelphia, with notes and descriptions of new species. Part 2», Proc. Acad. Nat. Sci. Phil., 12, 241 – 266.

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. Phil., 12[1860], 553 – 566.

Daudin F. M. (1803), Histoire naturelle générale et particulière des reptiles, Tome 6, F. Dufart, Paris.

Dolphin K., Belshaw R., Orme C. D. L., and Quicke D. L. J. (2000), «Noise and incongruence: interpreting results of the Incongruence Length Difference Test», Mol. Phylogen. Evol., 17(3), 401 – 406.

Dowling H. G. (1975a), «A provisional classification of snakes», in: Dowling H. G. (ed.), Yearbook of Herpetology, Vol. 1 [1974], HISS Publ., New York.

Dowling H. G. (1975b), «The Nearctic snake fauna», in: Dowling H. G. (ed.), Yearbook of Herpetology, Vol. 1 [1974], HISS Publ., New York.

Dowling H. G. and Duellmann W. E. (1978), «Subfamily Colubrinae», in: Systematic Herpetology: a Synopsis of Families and Higher Categories, HISS Publ. 7, New York.

Dowling H. G. and Fries I. (1987), «A taxonomic study of the ratsnakes. 8. A proposed new genus for Elaphe triaspis (Cope)», Herpetologica, 43(2), 200 – 207.

Dowling H. G., Hass C. A., Blair Hedges S., and Highton R. (1996), «Snake relationships revealed by slow-evolving proteins: a preliminary survey», J. Zool. Lond., 240(1), 1 – 28.

Dowling H. G., Highton R., Maha G. C., and Maxson L. R. (1983), «Biochemical evaluation of colubrid snake phylogeny», J. Zool. Lond., 201(3), 309 – 329.

Dowling H. G. and Maxson L. R. (1990), «Genetic and taxonomic relations of the short-tailed snakes, genus Stilosoma», J. Zool. Lond., 221(1), 77 – 85.

Dowling H. G. and Price R. M. (1988), «A proposed new genus for Elaphe subocularis and Elaphe rosaliae», The Snake, 20(1), 52 – 63.

Dugès A. (1897), «Description d’un Ophidien nouveau du Mexique (Oreophis boulengeri, g. et sp. nn.)», Proc. Zool. Soc. Lond., 1897(I), 284 – 285.

Duméril A. M. C., Bibron G., and Duméril A. (1854), Erpétologie générale ou histoire naturelle complète des reptiles, Vol. 7 (I), Roret, Paris.

Farris J. S. (1989), «The retention index and the rescaled consistency index», Cladistics, 5(4), 417 – 419.

Farris J. S., Kallersjö M., Kluge A. G., and Bult C. (1995), «Testing significance of incongruence», Cladistics, 10(3), 315 – 319.

Felsenstein J. (1985), «Confidence limits on phylogenies: an approach using the bootstrap», Evolution, 39(4), 783 – 791.

Fischer J. G. (1856), «Neue Schlangen des Hamburgischen Naturhistorischen Museums», Abh. Geb. Naturw. Hamb., 3, 81 – 116.

Fitzinger L. I. (1826), Neue Classification der Reptilien nach ihren natürlichen Verwandtschaften nebst einer Verwandtschafts-Tafel und einem Verzeichnisse der Reptiliensammlung des k. k. zoologischen Museum’s zu Wien, J. G. Heubner, Wien.

Fitzinger L. J. (1833), «Elaphe parreyssii — Parreyss’s Elaphe», in: Wagler J. (ed.), Descriptiones et Icones Amphibiorum (3), J. G. Cotta, München – Stuttgart – Tübingen.

Fitzinger L. (1843), Systema Reptilium. Amblyglossae, Braumüller et Seidel, Vindobonae [Vienna].

Gray J. E. (1853), «Descriptions of some undescribed species of reptiles collected by Dr. Joseph Hooker, in the Khasia Mountains, East Bengal and Sikkim, Himalaya», Ann. Mag. Nat. Hist. Ser. 2, 12, 386 – 392.

Griffin L. E. (1909), «A list of snakes found in Palawan», Phil. J. Sci., 4, 595 – 601.

Günther A. (1860), «Contribution to a knowledge of the reptiles of the Himalaya Mountains», Proc. Zool. Soc. Lond., 1860(I), 148 – 175.

Günther A. (1864a), The Reptiles of British India, R. Hardwicke [Ray Society], London.

Günther A. (1864b), «Report on a collection of reptiles and fishes from Palestine», Proc. Zool. Soc. Lond., 1864(III), 488 – 493.

Hallowell E. (1852), «Descriptions of new species of reptiles inhabiting North America», Proc. Acad. Nat. Sci. Phil., 6, 177 – 182.

Helfenberger N. (1989), Morphologie und Organtopographie bei Vertretern der Schlangengattung Elaphe Fitzinger, 1833 (Reptilia, Serpentes). Diploma Thesis, University of Zurich.

Helfenberger N. (2001a), Phylogenetic Relationships of Old World Ratsnakes Based on Visceral Organ Topography, Osteology, and Allozyme Variation. Ph.D. Thesis, University of Zurich.

Helfenberger N. (2001b), «Phylogenetic relationships of Old World Ratsnakes based on visceral organ topography, osteology, and allozyme variation», Russ. J. Herpetol., 8(Supplement), 1 – 62.

Holbrook J. E. (1842), North American Herpetology or a Description of the Reptiles Inhabiting the United States, Vol. 4 (2nd edition), J. Dobson, Philadelphia.

Holman A. J. (2000), Fossil Snakes of North America, Indiana Univ. Press, Bloomington.

Inger R. F. and Clark P. J. (1943), «Partition of the genus Coluber», Copeia, 1943(3), 141 – 145.

Jan G. (1863), «Enumerazione sistematica degli ofidi appartenenti al gruppo Coronellidae», Arch. Zool. Anat. Fis., 2(2), 213 – 330.

Jan G. (1865), «Prime linee d’una fauna della Persia occidentale», in: Note di un viaggio in Persia nel 1862 [F. De Filippi], G. Daelli et C. Editori, Milano.

Keogh J. S. (1996), «Evolution of the colubrid snake tribe Lampropeltini, a morphological perspective», Herpetologica, 52(3), 406 – 416.

Kramer E. (1977), «Zur Schlangenfauna Nepals», Rev. Suisse Zool., 84(3), 721 – 761.

Laurenti J. (1768), Austriaci Viennensis specimen medicum, exhibens synopsin reptilium, J. T. Trattnern, Wien.

Linnaeus C. (1758), Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Vol. 1 (editio decima, reformata), L. Salvii, Holmiae [Stockholm].

López T. J. and Maxson L. R. (1995), «Mitochondrial DNA sequence variation and genetic differentiation among colubrine snakes (Reptilia: Colubridae: Colubrinae)», Biochem. Syst. Ecol., 23(5), 487 – 505.

Lyons-Weiler J. (2001), RASA 3.0 Turbo for the Macintosh, http://bio.uml.edu/LW/RASA.html.

Lyons-Weiler J. and Hoelzer G. A. (1997), «Escaping from the Felsenstein zone by detecting long branches in phylogenetic data», Mol. Phylogen. Evol., 8(3), 375 – 384.

Lyons-Weiler J., Hoelzer G. A., and Tausch R. J. (1996), «Relative apparent synapomorphy analysis (RASA) I: the statistical measurement of phylogenetic signal», Mol. Phylogen. Evol., 13(6), 749 – 757.

Mattison C. (1999), Snake: The Essential Visual Guide to the World of Snakes, Dorling Kindersley, London.

Mertens R. and Rosenberg H. (1943), «Elaphe flavirufa (Cope), die mexikanische Nachtnatter», Wochenschr. Aquar. Terrar., 37(3), 60 – 62.

Michahelles [Dr.] (1833), «Rhinechis Agassizii — Agassiz’s Schnautzen-Natter», in: Wagler J. (ed.), Descriptiones et Icones Amphibiorum (3), J. G. Cotta, München – Stuttgart – Tübingen.

Nagy Z. T., Lawson R., Joger U., and Wink M. (2004), «Molecular systematics of racers, whipsnakes and relatives (Reptilia: Colubridae) using mitochondrial and nuclear markers», J. Zool. Syst. Evol. Res., 42(3), 223 – 233.

Nagy Z. T., Schmidtler J. F., Joger U., and Wink M. (2004), «Systematik der Zwergnattern (Reptilia: Colubridae: Eirenis) und verwandter Gruppen anhand von DNA-Sequenzen und morphologischen Daten», Salamandra, 39(3/4) [2003], 149 – 168.

Nei M. (1996), «Phylogenetic analysis in molecular evolutionary genetics», Ann. Rev. Gen., 30, 371 – 403.

Oldham J. C. and Smith H. M. (1991), «The generic status of the smooth green snake Opheodrys vernalis», Bull. Maryland Herpetol. Soc., 27(4), 201 – 215.

Oppel M. (1811), «Mémoire sur la classification des reptiles. Ordre II. Reptiles à écailles. Section II. Ophidiens (part 2)», Ann. Mus. Hist. Nat. Paris, 16(95)[1810], 376 – 393.

Ortenburger A. I. (1928), «The whipsnakes and racers, genera Masticophis and Coluber», Mem. Univ. Mich. Mus., 1, 1 – 247.

Peters W. (1862), «Eine Mittheilung von über die von dem so früh in Afrika verstorbenen Freiherrn von Barnim und Dr. Hartmann auf ihrer Reise durch Aegypten, Nubien und dem Sennâr gesammelten Amphibien», Monatsb. Kön. Pr. Akad. Wiss. Berl., 1862, 271 – 279.

Pinou T. and Dowling H. G. (2000), «’Coluber‘ versus ‘Ptyas‘: a regrettable tangle of names», Herpetol. Rev., 31(3), 136 – 138.

Pope C. H. (1935), Natural History of Central Asia. Vol. X. The Reptiles of China. Turtles, Crocodilians, Snakes, Lizards, Am. Mus. Nat. Hist., New York.

Posada D. and Crandall K. A. (1998), «MODELTEST: testing the model of DNA substitution», Bioinformatics, 14(9), 817 – 818.

Rzhetsky A. and Nei M. (1992), «A simple method for estimating and testing minimum evolution trees», Mol. Biol. Evol., 9(5), 945 – 967.

Rodríguez-Robles J. A. and De Jesús-Escobar J. M. (1999), «Molecular systematics of New World lampropeltinine snakes (Colubridae): implications for biogeography and evolution of food habits», Biol. J. Linn. Soc. Lond., 68(3), 355 – 385.

Saitou N. and Nei M. (1987), «The neighbor-joining method: A new method for reconstructing phylogenetic trees», Mol. Biol. Evol., 4(4), 406 – 425.

Sambrook J., Fritsch E., and Maniatis T. (1989), Molecular Cloning: A Laboratory Manual (2nd Edition), Vol. 1, Harbor Lab. Press, Cold Spring.

Schätti B. (1985), «Systematics of East African species of Meizodon Fischer, 1856 (Reptilia, Serpentes, Colubridae)», Monit. Zool. Ital. N. S., 20(8, Suppl.), 149 – 175.

Schätti B. (1986), «Morphological evidence for a partition of the genus Coluber (Reptilia: Serpentes)», in: Z. Roček (ed.), Studies in Herpetology. Proc. Eur. Herpetol. Meet., Prague, 1985, Charles University, Prague.

Schätti B. (1987), «The phylogenetic significance of morphological characters in the Holarctic racers of the genus Coluber», Amphibia–Reptilia, 8(4), 401 – 418.

Schätti B. (1988), Systematik und Evolution der Schlangengattung Hierophis Fitzinger, 1843 (Reptilia, Serpentes). Ph.D. Thesis, University of Zurich.

Schätti B. and Monsch P. (2004), «Systematics and phylogenetic relationships of Whip snakes (Hierophis Fitzinger) and Zamenis andreana Werner, 1917 (Reptilia: Squamata: Colubrinae)», Rev. Suisse Zool., 111(2), 239 – 256.

Schätti B. and Utiger U. (2001), «Hemerophis, a new genus for Zamenis socotrae Günther, and a contribution to the phylogeny of Old World racers, whip snakes and related genera (Reptilia: Squamata: Colubrinae)», Rev. Suisse Zool., 108(4), 919 – 948.

Schätti B. and Wilson L. D. (1986), Coluber Linnaeus — Holarctic racers (Reptilia: Squamata: Serpentes: Colubridae) [Cat. Am. Amph. Rept.], SSAR, Oxford.

Schlegel H. (1837), Essai sur la physionomie des serpens, Vol. 2. Partie descriptive, Lips et Van Stockum, La Haye [The Hague].

Schleich H. H. and Kästle W. (2002), Amphibians and Reptiles of Nepal [Biology, Systematics, Field Guide], Gantner, Ruggell.

Schmidt C. (2000), Morphologie der Hemipenes von Kletternattern (Elaphe Fitzinger, 1833). Diploma Thesis, University of Zurich.

Schmidt K. P. (1925), «New reptiles and a new salamander from China», Am. Mus. Novitates, 157, 1 – 5.

Schulz K.-D. (1996), A Monograph of the Colubrid Snakes of the Genus Elaphe Fitzinger, Koeltz Scientific Books, Havlicuv Brod.

Smith H. M. and Taylor E. H. (1945), «An annotated checklist and key to the snakes of Mexico», U.S. Nat. Mus. Bull., 187, IV + 1 – 239.

Smith M. A. (1943), The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese Sub-Region. Reptilia and Amphibia. Vol. III. Serpentes, Taylor et Francis, London.

Spolsky C. and Uzzell T. (1986), «Evolutionary history of the hybridogenetic hybrid frog Rana esculenta as deduced from mtDNA analyses», Mol. Biol. Evol., 3(1), 44 – 56.

Stejneger L. (1890), «On a new genus and species of colubrine snakes from North America», Proc. U.S. Nat. Mus., 13(802), 151 – 155.

Twerenbold P. (1987), Anatomische Untersuchungen an Vertretern der Schlangenfamilie Colubidae. Diploma Thesis, University of Zurich.

Underwood G. (1967), A Contribution to the Classification of Snakes, Trustees Br. Mus. (Nat. Hist.), London.

Utiger U., Helfenberger N., Schätti B., Schmidt C., Ruf M., and Ziswiler V. (2002), «Molecular systematics and phylogeny of Old and New World ratsnakes, Elaphe auct., and related genera (Reptilia, Squamata, Colubridae)», Russ. J. Herpetol., 9(2), 105 – 124.

Wagler J. G. (1828), «Gonyosoma viride — Grüner Kantenbauch», in: J. Wagler (ed.), Descriptiones et Icones Amphibiorum, J. G. Cotta, München.

Wagler J. (1830), Natürliches System der Amphibien mit vorangehender Classification der Säugethiere und Vögel, J. G. Cotta, München – Stuttgart – Tübingen.

Wall F. (1914), «Remarks to show that the snake hitherto known as Zamenis mucosus has been misplaced, and should be included in the genus Zaocys», J. Bombay Nat. Hist. Soc., 23(1), 168 – 169.

Wallach V. (1997), «A Monograph of the Colubrid Snakes of the Genus Elaphe Fitzinger, by Klaus-Dieter Schulz, 1996 [book review]», Herpetol. Rev., 28(2), 109 – 111.

Wallach V. (1998), «The lungs of snakes», in: Gans G. C. and Gaunt A. S. (eds.), Biology of the Reptilia. 19. Morphology C., SSAR, Ithaca.

Werner F. (1924), «Neue oder wenig bekannte Schlangen aus dem Naturhistorischen Staatsmuseum in Wien», Sitzungsb. Akad. Wiss. Wien. Math.-Naturw. Kl., [1]133(1), 29 – 59.

Whitaker R. and Captain A. (2004), Snakes of India, Draco Books, Chennai.

Williams K. L. and Wallach V. (1989), Snakes of the World, Vol. I. Synopsis of Snake Generic Names, Krieger, Malabar.




DOI: https://doi.org/10.30906/1026-2296-2005-12-1-39-60

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