PSEUDOXYRHOPHIINAE

Grazziotin, Hussam Zaher 1 Felipe Gobbi, Cadle, John E., Murphy, Robert W., Moura-Leite, Julio Cesar de & Bonatto, Sandro L., 2009, Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American Xenodontines: a revised classification and descriptions of new taxa, Papéis Avulsos de Zoologia 49 (11), pp. 115-153 : 137-139

SUBFAMILY PSEUDOXYRHOPHIINAE

Dowling, 1975 (Clade 18)

Pseudoxyrhophini Dowling, 1975.

Type-genus: Pseudoxyrhopus Günther, 1881 .

Diagnosis: (96%, 8). Spines reduced to spinules on the hemipenial lobes (Zaher, 1999).

Content: Alluaudina Mocquard, 1894 ; Amplorhinus A. Smith, 1847; Brygophis Domergue & Bour, 1989 ; Compsophis Mocquard, 1894 ; Ditypophis Günther, 1881 ; Dromicodryas Boulenger, 1893 ; Duberria Fitzinger, 1826 ; Exallodontophis Cadle, 1999 ; Heteroliodon Boettger, 1913 ; Ithycyphus Günther, 1873 ; Langaha Bonnaterre, 1790 ; Leioheterodon Jan, 1863 ; Liophidium Boulenger, 1896 ; Liopholidophis Mocquard, 1904 ; Lycodryas Günther, 1879 ; Madagascarophis Mertens, 1952 ; Micropisthodon Mocquard, 1894 ; Montaspis Bourquin 1991 ; Pararhadinaea Boettger, 1898 ; Pseudoxyrhopus Günther, 1881 ; Stenophis Boulenger, 1896 ; Thamnosophis Jan, 1863 .

Comments: The hemipenial synapomorphy of Pseudoxyrhophiinae is also present homoplastically in Homalopsidae . Geodipsas Boulenger, 1896 was placed in the synonymy of Compsophis by Glaw et al. (2007a). Bibilava Glaw, Nagy, Franzen & Vences, 2007 was synonymized with Thamnosophis ( Cadle & Ineich, 2008) . The broader phylogenetic analyses of Lawson et al. (2005) and Kelly et al. (2009) demonstrated convincingly that Duberria and Amplorhinus were more closely related to the Pseudoxyrhophiinae than to any other elapoid or colubroid lineage; a similar relationship of Amplorhinus (but not Duberria ) to pseudoxyrhophiids was previously suggested by Cadle (1994). Bourquin (1991) suggested, on the basis of skull morphology, that Montaspis is closely related to the Pseudoxyrhophiidae . We recognize both Stenophis and Lycodryas as valid, but the systematics of these snakes needs revision ( Cadle, 2003: 1000-1001); furthermore, Kelly et al. (2009) found that the two species of Stenophis they examined were not monophyletic relative to other pseudoxyrhophids. Species and generic level taxonomy of pseudoxyrhophids needs more research.

SUPERFAMILY COLUBROIDEA Oppel, 1811 (Clade 19)

Diagnosis: (98%, 10). Colubroids can be diagnosed by the presence of well-developed calyces present on the hemipenial lobes, a centrifugal sulcus spermaticus that divides on the proximal or central region of the hemipenial body and an aglyphous dentition.

Content: Calamariidae Bonaparte, 1838 ; Colubridae Oppel, 1811 ; Pseudoxenodontidae McDowell, 1987 ; Natricidae Bonaparte, 1838 ; Dipsadidae Bonaparte, 1838 .

Comments: Zaher (1999) discussed the variation regarding the sulcus spermaticus in colubroid snakes. Well-developed calyces on the hemipenial lobes are considered to be lost secondarily by the Natricidae . See above discussion on the new use of this name.

Family Calamariidae Bonaparte, 1838 (terminal taxon: Calamaria yunnanensis-pavimentata )

Calamarina Bonaparte, 1838: 392.

Type-genus: Calamaria H. Boie (in F. Boie), 1826.

Diagnosis: Frontals and sphenoid forming ventral border of the optic foramen (excluding entirely, or nearly so, the parietals); hemipenial body nude; hemipenial body bearing a pair of longitudinal ridges (Zaher, 1999).

Content: Calamaria H. Boie (in F. Boie), 1826; Calamorhabdium Boettger, 1898 ; Collorhabdium Smedley, 1932 ; Etheridgeum Wallach, 1988 ; Macrocalamus Günther, 1864 ; Pseudorabdion Jan, 1862 ; Rabdion Duméril, 1853 .

FAMILY COLUBRIDAE Oppel, 1811 (Clade 21)

Colubrini Oppel, 1811:50.

Type-genus: Coluber Linnaeus, 1758 .

Diagnosis: (97%, 7). Sulcus spermaticus simple, derived from the right branch of a primitively divided sulcus (see Comments).

Content: Aeluroglena Boulenger, 1898 ; Ahaetulla Link, 1807 ; Argyrogena Werner, 1924 ; Arizona Kennicott (in Baird), 1859; Bogertophis Dowling and Price, 1988 ; Boiga Fitzinger, 1826 ; Cemophora Cope, 1860 ; Chilomeniscus Cope, 1860 ; Chionactis Cope, 1860 ; Chironius Fitzinger, 1826 ; Chrysopelea H. Boie (in Schlegel), 1826; Coelognathus Fitzinger, 1843 ; Coluber Linnaeus, 1758 ; Conopsis Günther, 1858 ; Coronella Laurenti, 1768 ; Crotaphopeltis Fitzinger, 1843 ; Cryptophidion Wallach and Jon 1992 ; Cyclophiops Boulenger, 1888 ; Dasypeltis Wagler, 1830 ; Dendrelaphis Boulenger, 1890 ; Dendrophidion Fitzinger, 1843 ; Dinodon Duméril, Bibron & Duméril, 1854 ; Dipsadoboa Günther, 1858 ; Dispholidus Duvernoy, 1832 ; Drymarchon Fitzinger, 1843 ; Drymobius Fitzinger, 1843 ; Drymoluber Amaral, 1930 ; Dryocalamus Günther, 1858 ; Dryophiops Boulenger, 1896 ; Eirenis Jan, 1863 ; Elachistodon Reinhardt, 1863 ; Elaphe Fitzinger (in Wagler), 1833; Euprepiophis Fitzinger, 1843 ; Ficimia Gray, 1849 ; Gastropyxis Cope, 1861 ; Geagras Cope, 1875 ; Gonyophis Boulenger, 1891 ; Gonyosoma Wagler, 1828 ; Gyalopion Cope, 1860 ; Hapsidophrys Fischer, 1856 ; Hemerophis Schätti & Utiger, 2001 ; Hemorrhois F. Boie, 1826 ; Hierophis Fitzinger (in Bonaparte), 1834; Lampropeltis Fitzinger, 1843 ; Leptodrymus Amaral, 1927 ; Leptophis Bell, 1825 ; Lepturophis Boulenger, 1900 ; Liochlorophis Oldham & Smith, 1991 ; Liopeltis Fitzinger, 1843 ; Lycodon Boie (in Fitzinger), 1826; Lytorhynchus Peters, 1862 ; Macroprotodon Duméril & Bibron (in Guichenot), 1850; Maculophis Burbrink & Lawson, 1997; Masticophis Baird (in Baird & Girard), 1853; Mastigodryas Amaral, 1934 ; Meizodon Fischer, 1856 ; Oligodon H. Boie (in Fitzinger), 1826; Oocatochus Helfenberger, 2001 ; Opheodrys Fitzinger, 1843 ; Oreocryptophis Utiger, Schätti & Helfenberger, 2005 ; Oreophis Utiger, Helfenberger, Schätti, Schmidt, Ruf & Ziswiler, 2002 ; Orthriophis Utiger, Helfenberger, Schätti, Schmidt, Ruf & Ziswiler, 2002 ; Oxybelis Wagler, 1830 ; Pantherophis Fitzinger, 1843 ; Philothamnus A. Smith, 1847; Phyllorhynchus Stejneger, 1890 ; Pituophis Holbrook, 1842 ; Platyceps Blyth, 1860 ; Pseudelaphe Mertens & Rosenberg, 1943 ; Pseudocyclophis Boettger, 1888 ; Pseudoficimia Bocourt, 1883 ; Pseustes Fitzinger, 1843 ; Ptyas Fitzinger, 1843 ; Rhamnophis Günther, 1862 ; Rhinechis Michahelles, 1833 ; Rhinobothryum Wagler, 1830 ; Rhinocheilus Girard (in Baird & Girard), 1853; Rhynchocalamus Günther, 1864 ; Rhynchophis Mocquard, 1897 ; Salvadora Baird (in Baird & Girard), 1853; Scaphiodontophis Taylor & Smith, 1943 ; Scaphiophis Peters, 1870 ; Scolecophis Fitzinger, 1843 ; Senticolis Dowling & Fries, 1987 ; Sibynophis Fitzinger, 1843 ; Simophis Peters, 1860 ; Sonora Girard (in Baird & Girard), 1853; Spalerosophis Jan (in De Filippi), 1865 ; Spilotes Wagler, 1830 ; Stegonotus Duméril, Bibron & Duméril, 1854 ; Stenorrhina Duméril, 1853 ; Stilosoma Brown, 1890 ; Symphimus Cope, 1870 ; Sympholis Cope, 1862 ; Tantilla Girard (in Baird & Girard), 1853; Tantillita Smith, 1941 ; Telescopus Wagler, 1830 ; Thelotornis A. Smith, 1849; Thrasops Hallowell, 1857 ; Toxicodryas Hallowell 1857 ; Trimorphodon Cope, 1861 ; Xenelaphis Günther, 1864 ; Xyelodontophis Broadley & Wallach 2002 ; Zamenis Bonaparte, 1838 ; Zaocys Cope, 1861 .

Comments: Use of the name “ Colubridae ” for this clade is a much more restricted use of this name than its long-standing use in the literature on caenophidian systematics, in which “ Colubridae ” generally referred to all caenophidians that were not acrochordids, elapids, or viperids. The single sulcus spermaticus of colubrids and natricids is considered to have derived from a centrifugally divided sulcus, but in different ways in the two groups ( McDowell 1961). On unilobed organs of colubrids the sulcus extends centrolineally to the distal end of the hemipenis, whereas on some distally bilobed organs the sulcus always extends to the right lobe. On the other hand, in natricids when the sulcus extends to only one of the lobes of a bilobed organ, it is always to the left lobe (see also Rossman & Eberle, 1977; and Zaher, 1999: 25-26). Lawson et al. (2005) have shown that Macroprotodon lies within the family Colubridae , but without clear affinities within that group. The phylogenetic affinities of Scaphiophis Peters, 1870 has been disputed (Zaher, 1999; Vidal et al., 2008). Recently, Kelly et al. (2008) included the genus in their molecular analysis, in which it appears nested within colubrines. For this reason, we include this genus in the family Colubridae .