SSAR Common Names Checklist ver. 2022-02-11

anura — frogs

a. blanchardi Blanchard's Cricket Frog

Gamble et al. (2008, Mol. Phylogenet. Evol. 48: 112–125) recognized Acris blanchardi as distinct from A. crepitans on the basis of molecular evidence (and included Acris crepitans paludicola as a synonym of A. blanchardi), although McCallum and Trauth (2006, Zootaxa 1104: 1–21) previously rejected the distinctiveness of A. c. blanchardi from A. c. crepitans on the basis of morphology. Reviewed by Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press. 205–219).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-20

a. crepitans Eastern Cricket Frog

See comment under Acris blanchardi. Reviewed by Gray et al. (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ of California Press: 441–443), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press. 219–226).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-20

a. gryllus Southern Cricket Frog

The lineages delimited on the basis of the molecular evidence of Gamble et al. (2008, Mol. Phylogenet. Evol. 48: 112-125) do not correspond to the nominal subspecies occasionally employed by various previous authors. It seems on that basis that recognition of the subspecies. A. g. dorsalis and A. g. gryllus, is not warranted. Reviewed by Jensen (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ of California Press: 443–444).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-20

a. americanus American Toad

Geographic variation has been insu ciently studied, although careful evaluation of call and/or molecular data might provide considerable evidence of divergent lineages. See comments under A. baxteri, A. fowleri, A. hemiophrys, A. terrestris, and A. woodhousii. Masta et al. (2002, Mol. Phylogenet. Evol. 24: 302–314) provided evidence that suggests that A. a. charlesmithi may be a distinct species. Reviewed by Green (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 692–704) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press. 219–226).

Note on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-20

a. americanus americanus Eastern American Toad

a. americanus charlesmithi Dwarf American Toad

a. baxteri Wyoming Toad

Recognized as a species, rather than a subspecies of A. hemiophrys by Packard (1971, J. Herpetol. 5: 191–193), and more recently by Smith et al. (1998, Contemp. Herpetol. 1). Nevertheless, Cook (1983, Publ. Nat. Sci. Natl. Mus. Canada 3) considered A. baxteri to be undiagnosable against the background of geographic variation in A. hemiophrys (as Bufo americanus hemiophrys), and this has not been addressed by subsequent authors. Reviewed by Odum and Corn (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press:390–392), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press. 43–47).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-20

a. boreas Western Toad

See Schuierer (1963, Herpetologica 18: 262–267). Two nominal subspecies are generally recognized, although Goebel (2005, In Lannoo, M. [ed.], Amphibian Declines, Univ. California Press, pp. 210–211) discussed geographic variation and phylogenetics of the A. boreas (as the Bufo boreas) group (i.e., A. boreas, A. canorus, A. exsul, and A. nelsoni), and noted other unnamed populations of nominal A. boreas that may be species. Populations in Alberta, Canada, assigned to A. boreas have a distinct breeding call and vocal sacs (Cook, 1983, Publ. Nat. Sci. Natl. Mus. Canada 3; Pauly 2008, PhD Dissertation, Univ. Texas at Austin); the taxonomic implications of this warrant investigation. Goebel et al. (2009, Mol. Phylogenet. Evol. 50: 209–225) suggested on the basis of molecular evidence that nominal Anaxyrus boreas is a complex of species (as suggested previously by Bogert, 1960, Animal Sounds Commun: 179) that do not conform to the traditional limits of taxonomic species and subspecies (and which we do not recognize here for this reason) and that some populations assigned to this taxon may actually be more closely related to Anaxyrus canorus and A. nelsoni—a problem that calls for additional elucidation. Reviewed by Muths and Nanjappa (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 392–396) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 47–65).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-20

a. californicus Arroyo Toad

See Gergus (1998, Herpetologica 54: 317–325) for justification for this to be considered a distinct species from Anaxyrus microscaphus. Reviewed by Price and Sullivan (1988, Cat. Am. Amph. Rept. 415, as Bufo microscaphus californicus), Sweet and Sullivan (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 396–400), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 65–70).

Note on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

a. canorus Yosemite Toad

Reviewed by Karlstrom (1973, Cat. Am. Amph. Rept. 132), Davidson and Fellers (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 400–401), and Dodd (2013, Frogs U.S. and Canada, 1: 70–77). See comment under A. boreas.

Note on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

a. cognatus Great Plains Toad

Reviewed by Krupa (1990, Cat. Am. Amph. Rept. 457), Graves and Krupa (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 440–404) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 78–87).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-28

a. debilis Chihuahuan Green Toad

See accounts in Sanders and Smith (1951, Field and Laboratory 19: 141–160) and by Bogert (1962, Am. Mus. Novit. 2100) as Bufo debilis. Reviewed by Painter (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 404–406, as Bufo debilis) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 88–91). The nominal subspecies are unlikely to be anything other than arbitrarily defined sections of clines although this remains to be investigated adequately. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation: 301) rejected subspecies but presented no evidence for this conclusion.

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-28

a. debilis debilis Eastern Chihuahuan Green Toad

See accounts in Sanders and Smith (1951, Field and Laboratory 19: 141–160) and by Bogert (1962, Am. Mus. Novit. 2100) as Bufo debilis. Reviewed by Painter (2005, in Lannoo, M. [ed.], Amph. Declines: 404–406, as Bufo debilis) and Dodd (2013, Frogs U.S. and Canada, 1: 88–91). The nominal subspecies are unlikely to be anything other than arbitrarily defined sections of clines although this remains to be investigated adequately. Fouquette and Dubois (2014, Checklist N.A. Amph. Rept.: 301) rejected subspecies but presented no evidence for this conclusion.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

a. debilis insidior Western Chihuahuan Green Toad

See accounts in Sanders and Smith (1951, Field and Laboratory 19: 141–160) and by Bogert (1962, Am. Mus. Novit. 2100) as Bufo debilis. Reviewed by Painter (2005, in Lannoo, M. [ed.], Amph. Declines: 404–406, as Bufo debilis) and Dodd (2013, Frogs U.S. and Canada, 1: 88–91). The nominal subspecies are unlikely to be anything other than arbitrarily defined sections of clines although this remains to be investigated adequately. Fouquette and Dubois (2014, Checklist N.A. Amph. Rept.: 301) rejected subspecies but presented no evidence for this conclusion.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

a. exsul Black Toad

See comment under A. boreas. Reviewed by Fellers (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 406–408, as Bufo exsul) and Dodd (2013, Frogs U.S. and Canada, 1: 92–96).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-28

a. fowleri Fowler's Toad

Green (1996, Israel J. Zool. 42: 95–109) discussed the problem of interspecific hybridization in the A. americanus complex and briefly addressed the publication by Sanders (1987, Evolutionary hybridization and speciation in North American indigenous bufonids. O. Sanders, Dallas, TX), in which Sanders recognized a number of dubiously delimited taxa within the A. americanus complex (his Bufo hobarti, which would be in the synonymy of A. fowleri; Bufo copei, which would be in A. americanus, and Bufo planiorum and Bufo antecessor, both of which would be in the synonymy of A. woodhousii woodhousii). None have been formally synonymized, nor have any attracted recognition by those working on the complex. See comment under A. woodhousii. Masta et al. (2002, Mol. Phylogenet. Evol. 24: 302–314) provided evidence for the distinctiveness of this species from A. woodhousii and noted (as did Smith and Green, 2004, Mol. Ecol. 13: 3723–3733) that at the molecular level there are multiple, distinct mitochondrially-recognizable populations in A. fowleri. Reviewed by Green (2005, in Lannoo, M. J. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press:, as Bufo fowleri) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 96–113).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-28

a. hemiophrys Canadian Toad

See comment under A. baxteri. Cook (1983, Publ. Nat. Sci. Natl. Mus. Canada 3) regarded A. hemiophrys and A. americanus as forming very distinctive subspecies of one species, although subsequent authors (e.g., Green and Pustowka, 1997, Herpetologica 53: 218–228) have regarded the contact zone between these taxa as a hybrid zone between two species. Reviewed by Ewert and Lannoo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press:412–415, as Bufo hemiophrys) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 113–120).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-28

a. houstonensis Houston Toad

Reviewed by Brown (1973, Cat. Am. Amph. Rept. 133, as Bufo houstonensis), Shepard and Brown (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press:: 415–417, as Bufo houstonensis), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 120–126).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-28

a. microscaphus Arizona Toad

Reviewed by Price and Sullivan (1988, Cat. Am. Amph. Rept. 415, as Bufo microscaphus), Schwaner and Sullivan (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 422–424, as Bufo microscaphus), and Dodd, 2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 127–13). See comment under A. californicus. Formerly included A. californicus and A. mexicanus (extralimital) as subspecies, both of which were recognized as species by Gergus (1998, Herpetologica 54: 317–325).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. nelsoni Amargosa Toad

Stebbins (1985, A Field Guide to Western Reptiles and Amphibians, Houghton Mifflin, Boston) and Altig et al. (1998, Contemp. Herpetol. Inform. Serv. 2) regarded A. nelsoni as a species, rather than a subspecies of A. boreas. Reviewed by Goebel, Smith, Murphy, and Morafka (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 427–430, as Bufo nelsoni) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 132–136). See comment under A. boreas.

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. punctatus Red-Spotted Toad

Reviewed by Korky (1999, Cat. Am. Amph. Rept. 1104, as Bufo punctatus), Sullivan (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 430–432, as Bufo punctatus), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 136–144).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

a. quercicus Oak Toad

Reviewed by Ashton and Franz (1979, Cat. Am. Amph. Rept. 222, as Bufo quercicus), Punzo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 432–433, as Bufo quercicus), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 144–149).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. retiformis Sonoran Green Toad

Reviewed by Hulse (1978, Cat. Am. Amph. Rept. 207, as Bufo retiformis), Blomquist (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 433–435, as Bufo retiformis), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 149–152).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. speciosus Texas Toad

Older literature confused this species with A. cognatus, A. mexicanus (extralimital), and A. compactilis (extralimital). Rogers (1972, Copeia 1972: 381–383) demonstrated its morphological distinctiveness. Reviewed by Dayton and Painter (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 435–436, as Bufo speciosus), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 152–155).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. terrestris Southern Toad

No reports of geographic variation exist in the literature, although extensive geographic variation is evident on examination of specimens. Hybridization with A. americanus along the Fall Line may have strong effects on geographic variation, although data on this have not been published. Reviewed by Blem (1979, Cat. Am. Amph. Rept. 223, as Bufo terrestris), Jensen (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 436–438, as Bufo terrestris), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 155–166).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. woodhousii Woodhouse's Toad

See comments under A. fowleri. The incorrect spelling of the species name to woodhousei has been used widely. The status of taxa recognized by Sanders (1987, Evol. Hybrid. Spec. N. Am. Indig. Bufonids) has not been evaluated closely by any author, although neither have they enjoyed any recognition. Evidence provided by Masta et al. (2002, Mol. Phylogenet. Evol. 24: 302–314) suggests that A. w. australis may be a distinct species and that former A. w. velatus is a hybrid population of A. woodhousii × A. fowleri, and therefore should not be recognized. Reviewed by Sullivan (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 438–440, as Bufo woodhousii) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 166–176).

Notes on genus:
This taxon of strictly North American toads was removed from “Bufo” (as well as were a number of other taxa) by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) as a revision to render a monophyletic taxonomy and with genera delimited to be more compact than the unwieldy "Bufo". The phylogenetic study of bufonids by Van Bocxlaer et al. (2010, Science 327: 679–682) also suggests that New World "Bufo" do not form a monophyletic group. Smith and Chiszar (2006, Herpetol. Conserv. Biol. 1: 6-8) recommend retaining the North American taxa Anaxyrus, Incilius, and Rhinella (as well as such long-recognized extralimital taxa such as Ansonia, Capensibufo, Crepidophryne, Didynamipus, Mertensophryne, Nectophryne, Nectophrynoides, Pedostibes, Pelophryne, Schismaderma, Werneria, and Wolterstorffina) as subgenera of Bufo to obviate the need for generic changes in North American species. More recently, Fouquette and Dubois (2014, Checklist N. Am. Amph. Rept.) followed this approach in a modified form. This approach, though, would visit considerable nomenclatural instability on many countries outside of the USA and Canada. See Pauly et al. (2009, Herpetologica 65: 115–128) and Frost et al. (Herpetologica 65: 136–153) for discussion.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. woodhousii australis Southwestern Woodhouse's Toad

a. woodhousii woodhousii Rocky Mountain Toad

a. montanus Rocky Mountain Tailed Frog

See Nelson et al. (2001, Evolution 55: 147–160) for evidence supporting the recognition of this species distinct from A. truei. Adams (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 382) provided a brief but detailed review as did Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 17).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

a. truei Coastal Tailed Frog

See Metter (1968, Cat. Am. Amph. Rept. 69) for review (as including A. montanus). Reviewed by Adams and Pearl (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 382–385) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 7¬–16).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-29

c. augusti Barking Frog

Reviewed by Zweifel (1967, Cat. Am. Amph. Rept. 41, as Eleutherodactylus augusti) and Schwalbe and Goldberg, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 491–492). Goldberg et al. (2004, Herpetologica 60: 312–320) suggested that C. a. cactorum and C. a. latrans are different species but did not provide a formal new taxonomy.

Note on genus:
This taxon of predominantly Mexican and Central American frogs was removed from a paraphyletic “Eleutherodactylus” by Crawford and Smith (2005, Mol. Phylogenet. Evol. 35: 536–555).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

c. augusti cactorum Western Barking Frog

Note on genus:
This taxon of predominantly Mexican and Central American frogs was removed from a paraphyletic “Eleutherodactylus” by Crawford and Smith (2005, Mol. Phylogenet. Evol. 35: 536–555).

Reviewed by Zweifel (1967, Cat. Am. Amph. Rept. 41, as Eleutherodactylus augusti) and Schwalbe and Goldberg, I2005, in Lannoo, M. [ed.], Amph. Declines: 491–492). Goldberg et al. (2004, Herpetologica 60: 312–320) suggested that C. a. cactorum and C. a. latrans are different species but did not provide a formal new taxonomy.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

c. augusti latrans Balcones Barking Frog

Note on genus:
This taxon of predominantly Mexican and Central American frogs was removed from a paraphyletic “Eleutherodactylus” by Crawford and Smith (2005, Mol. Phylogenet. Evol. 35: 536–555).

Reviewed by Zweifel (1967, Cat. Am. Amph. Rept. 41, as Eleutherodactylus augusti) and Schwalbe and Goldberg, I2005, in Lannoo, M. [ed.], Amph. Declines: 491–492). Goldberg et al. (2004, Herpetologica 60: 312–320) suggested that C. a. cactorum and C. a. latrans are different species but did not provide a formal new taxonomy.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

d. auratus Green-And-Black Poison Dart Frog

Alien Species:

The most recent review of this genus and its relatives is Grant et al. (2006, Bull. Amer. Mus. Nat. Hist. 299: 1–262).

The Green-and-black Poison Dart Frog is native to Central America and Colombia and is established in Hawaii.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

e. coqui Coquí

Alien Species:

The Coquí is native to Puerto Rico, has been reported from five states, is established in Hawaii, and is established in a few greenhouses in California. It is widely established on Hawaii Island but is more restricted and the target of eradication efforts on the other Hawaiian Islands.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

e. cystignathoides Rio Grande Chirping Frog

Note on genus:
See Craugastor. Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) recognized Syrrhophus for a monophyletic group containing E. cystignathoides, E. guttilatus, and E. marnocki and Euhyas for a group containing E. planirostris. Heinicke et al. (2007, Proc. Natl. Acad. Sci. USA 104: 10092–97) and Hedges et al. (2008, Zootaxa 1737: 1-182) redelimited Eleutherodactylus as monophyletic by exclusion of a number of South American taxa and treated (and redelimited) Euhyas and Syrrhophus as subgenera of Eleutherodactylus.

Two nominal subspecies named, of which only one of which enters the USA. The status of these taxa, whether they represent arbitrarily delimited parts of a single population or different lineages is unknown. Reviewed by Wallace, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 494–495) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 197–199).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

e. cystignathoides campi Rio Grande Chirping Frog

Note on genus:
See Craugastor. Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) recognized Syrrhophus for a monophyletic group containing E. cystignathoides, E. guttilatus, and E. marnocki and Euhyas for a group containing E. planirostris. Heinicke et al. (2007, Proc. Natl. Acad. Sci. USA 104: 10092–97) and Hedges et al. (2008, Zootaxa 1737: 1-182) redelimited Eleutherodactylus as monophyletic by exclusion of a number of South American taxa and treated (and redelimited) Euhyas and Syrrhophus as subgenera of Eleutherodactylus.

Two nominal subspecies named, of which only one of which enters the USA. The status of these taxa, whether they represent arbitrarily delimited parts of a single population or different lineages is unknown. Reviewed by Wallace, (2005, in Lannoo, M. [ed.], Amph. Declines: 494–495) and Dodd (2013, Frogs U.S. and Canada, 1: 197–199).

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

e. guttilatus Spotted Chirping Frog

Geographic variation is poorly known. Some authors (e.g. Morafka, 1977, Biogeographica 9) considered E. guttilatus to be a synonym of E. c. campi (and by extension, of E. cystignathoides) but this remains to be sufficiently tested. Reviewed by Wallace (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 496–497) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 199–201).

Note on genus:
See Craugastor. Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) recognized Syrrhophus for a monophyletic group containing E. cystignathoides, E. guttilatus, and E. marnocki and Euhyas for a group containing E. planirostris. Heinicke et al. (2007, Proc. Natl. Acad. Sci. USA 104: 10092–97) and Hedges et al. (2008, Zootaxa 1737: 1-182) redelimited Eleutherodactylus as monophyletic by exclusion of a number of South American taxa and treated (and redelimited) Euhyas and Syrrhophus as subgenera of Eleutherodactylus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

e. marnockii Cliff Chirping Frog

See account by Lynch (1970, Univ. Kansas Publ. Mus. Nat. Hist. 20: 1–45) and reviews by Wallace, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 201–204). Geographic variation is not well studied.

Note on genus:
See Craugastor. Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297) recognized Syrrhophus for a monophyletic group containing E. cystignathoides, E. guttilatus, and E. marnocki and Euhyas for a group containing E. planirostris. Heinicke et al. (2007, Proc. Natl. Acad. Sci. USA 104: 10092–97) and Hedges et al. (2008, Zootaxa 1737: 1-182) redelimited Eleutherodactylus as monophyletic by exclusion of a number of South American taxa and treated (and redelimited) Euhyas and Syrrhophus as subgenera of Eleutherodactylus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

e. planirostris Greenhouse Frog

Alien Species:

The Greenhouse Frog is native to Cuba, The Bahamas, and Cayman Islands and is established in Alabama, Florida, Georgia, Hawaii, Louisiana, Mississippi, and South Carolina.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

g. carolinensis Eastern Narrow-Mouthed Toad

Reviewed by Nelson (1972, Cat. Am. Amph. Rept. 120), Mitchell and Lannoo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 501–503) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 439–448).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

g. mazatlanensis Sinaloan Narrow-Mouthed Toad

Recognized as distinct from G. olivacea by Streicher et al. (2012, Mol. Phylogenet. Evol. 64: 645-653).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

g. olivacea Western Narrow-Mouthed Toad

Reviewed by Nelson (1972, Cat. Am. Amph. Rept. 122), Sredl and Field (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 503–506), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 448–455) in the sense of including G. mazatlanensis of southern Arizona.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

g. rugosa Japanese Wrinkled Frog

Alien Species:

This genus of Asian frogs was recently removed from a polyphyletic “Rana” by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297).

The Japanese Wrinkled Frog is native to Japan and is established in Hawaii.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

h. andersonii Pine Barrens Treefrog

Reviewed by Gosner and Black (1967, Cat. Am. Amph. Rept. 54), Means (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 445–447), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 235–239). The widely disjunct populations have been examined with allozymes and only subtle (no fixed differences) geographic variation was documented (Karlin et al., 1982, Copeia 1982: 175–178).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. arenicolor Canyon Treefrog

Barber (1999, Mol. Ecol. 8: 563–576) examined geographic variation and suggested that at least two other species should be recognized within the Mexican component of its range. Bryson et al. (2010, Evolution, 64: 2315-2340) also reported on molecular geographic variation and demonstrated introgression with Hyla wrightorum. Reviewed by Painter (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 447–448) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 239–245).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. avivoca Bird-Voiced Treefrog

Smith (1953, Herpetologica 9: 169–173) discussed geographic variation and recognized two nominal subspecies which are rarely employed. Reviewed by Smith (1966, Cat. Am. Rept. Amph. 28), Redmer (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 448–449) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 245–250).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. avivoca avivoca Western Bird-Voiced Treefrog

Smith (1953, Herpetologica 9: 169–173) discussed geographic variation and recognized two nominal subspecies which are rarely employed. Reviewed by Smith (1966, Cat. Am. Rept. Amph. 28), Redmer (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 448–449) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 245–250).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. avivoca ogechiensis Eastern Bird-Voiced Treefrog

Smith (1953, Herpetologica 9: 169–173) discussed geographic variation and recognized two nominal subspecies which are rarely employed. Reviewed by Smith (1966, Cat. Am. Rept. Amph. 28), Redmer (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 448–449) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 245–250).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. chrysoscelis Cope's Gray Treefrog

See comment under H. versicolor. Reviewed by Hoffman (1988, Cat. Am. Amph. Rept. 436), Cline, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 449–452), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 250–262).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. cinerea Green Treefrog

Subspecies occasionally are recognized (H. c. cinerea and H. c. evittata) without discussion, and on the basis of a single populationally variable character. See Duellman and Schwartz (1958, Bull. Florida State Mus., Biol. Sci. 3: 241) for discussion and rejection of subspecies. Reviewed by Redmer and Brandon (2003, Cat. Am. Amph. Rept. 766), Redmer and Brandon (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 452–454), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 262–273).

Note on genus:
Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. femoralis Pine Woods Treefrog

Reviewed by Hoffman (1988, Cat. Am. Amph. Rept. 436), Mitchell (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 454–456), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 274–280).

Notes on genus: Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. gratiosa Barking Treefrog

Reviewed by Caldwell (1982, Cat. Am. Amph. Rept. 298), Mitchell (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 455–456), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 280–288).

Notes on genus: Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. squirella Squirrel Treefrog

Reviewed by Martof (1975, Cat. Am. Amph. Rept. 168), Mitchell and Lannoo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 456–458), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 288–294).

Notes on genus: Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. versicolor Gray Treefrog

Holloway et al. (2006, Am. Nat. 167: E88–E101) discussed the role of diploid H. chrysoscelis in the formation of the tetraploid H. versicolor, reviewed previous literature, and provided a revised range. Reviewed by Cline, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 458–461) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 294–309).

Notes on genus: Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. wrightorum Arizona Treefrog

Gergus et al. (2004, Copeia 2004: 758–769) reported on the distinctiveness of this species with respect to H. eximia (extralimital). See comment under H. arenicolor. Reviewed by Gergus, Wallace, and Sullivan (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 461–463) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 309–332).

Notes on genus: Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist., 294) redelimited this monophyletic taxon to include only North American and Eurasian species. Hua et al. (2009, Herpetologica 65: 246–259) discussed relationships within the group. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation) recently recognized a suite of subgenera based on genetic and morphological evidence, but pending a more thorough evidentiary review, we hesitate to employ this taxonomy. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted Hyla to Eurasia and North Africa and referred the North American and east Asian sister taxon of this group to Dryophytes, although the acceptance of this taxonomy within the community is not clear at this point.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

h. variolosus Sheep Frog

See Nelson (1973, Herpetologica 29: 6–17; 1974, Herpetologica 30: 250–274) for discussion of geographic variation and rejection of subspecies. USA population reviewed by Judd and Irwin (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 506–508) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 455–457). Although only two species are currently recognized within this genus, very strong geographic variation in coloration, call, and toe structure suggests that several species are masquerading under this particular name. Given that the type locality of H. variolosus is in Costa Rica, the scientific name applied to the U.S. form is likely to change.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

i. alvarius Sonoran Desert Toad

Reviewed by Fouquette (1970, Cat. Am. Amph. Rept. 93, as Bufo alvarius), Fouquette et al. (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 384–386, as Bufo alvarius), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 177–180).

Note on genus:
This taxon of predominantly Central American toads was removed from a paraphyletic “Bufo” by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297; as Cranopsis). However, the oldest name for this taxon is Incilius Cope, 1863 (see Frost et al., 2009, Copeia 2009: 418–419) which therefore takes precedence. See comment under Anaxyrus. Van Bocxlaer et al. (2010, Science, 327: 679–682) presented evidence that Incilius may be paraphyletic with respect to Anaxyrus due to the placement of one extralimital species, although this was based on a small dataset (Mendelson et al., 2011, Zootaxa, 3138: 1-34). See comment under Anaxyrus, regarding the treatment of this genus as a subgenus of Bufo by some although the effect extralimitally of subgeneric status would be to require a number well-marked genera (e.g., Ansonia) to be treated as subgenera as well.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

i. nebulifer Gulf Coast Toad

Mulcahy and Mendelson (2000, Mol. Phylogenet. Evol. 17: 173) recognized this species, as Bufo nebulifer, and as distinct from I. valliceps, an extralimital species. Reviewed by Mendelson (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 424–427, as Bufo nebulifer), Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 180–186), and Mendelson et al. (2015, Zootaxa 3974: 517–537).

Note on genus:
This taxon of predominantly Central American toads was removed from a paraphyletic “Bufo” by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297; as Cranopsis). However, the oldest name for this taxon is Incilius Cope, 1863 (see Frost et al., 2009, Copeia 2009: 418–419) which therefore takes precedence. See comment under Anaxyrus. Van Bocxlaer et al. (2010, Science, 327: 679–682) presented evidence that Incilius may be paraphyletic with respect to Anaxyrus due to the placement of one extralimital species, although this was based on a small dataset (Mendelson et al., 2011, Zootaxa, 3138: 1-34). See comment under Anaxyrus, regarding the treatment of this genus as a subgenus of Bufo by some although the effect extralimitally of subgeneric status would be to require a number well-marked genera (e.g., Ansonia) to be treated as subgenera as well.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. fragilis Mexican White-Lipped Frog

Reviewed by Heyer et al. (2006, Cat. Am. Amph. Rept. 830), Heyer (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 500–501), Dodd, 2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 436–438. Much of the older literature about this species refers to it incorrectly as Leptodactylus labialis.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. areolatus Crawfish Frog

See comment under L. capito. Reviewed by Altig and Lohoefener (1983, Cat. Am. Amph. Rept. 324, as Rana areolata), Parris and Redmer (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 526–528), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 461–466). Geographic variation deserves further study to determine status of the nominal subspecies.

Notes on genus: This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. areolatus areolatus Southern Crawfish Frog

Notes on genus: This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. areolatus circulosus Northern Crawfish Frog

Notes on genus: This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. berlandieri Rio Grande Leopard Frog

Geographic variation is not well documented and relationships with extralimital Mexican forms (e.g., L. forreri, L. brownorum) are not well understood. Reviewed with special reference to the USA populations by Rorabaugh (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 530–532) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 466–471).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. blairi Plains Leopard Frog

Reviewed by Brown (1992, Cat. Am. Amph. Rept. 536, as Rana blairi) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 472–479). Isolated western populations have not been well studied.

Notes on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. capito Gopher Frog

Lithobates capito is considered by some to be part of L. areolatus (but see Case, 1978, Syst. Zool. 27: 299–311, who considered them distinct). Reviewed by Altig and Lohoefener (1983, Cat. Am. Amph. Rept. 324, as Rana areolata capito), Jensen and Richter (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 536–538), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 479–485). Recognized as distinct from L. areolatus by Young and Crother (2001, Copeia, 2001: 382–388), who also rejected subspecies. Richter et al. (2014, Copeia: 231–237) presented mitochondrial evidence on interpopulational variation at the molecular level and suggested an historical structure among these.

Notes on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. catesbeianus American Bullfrog

Geographic variation within the natural range L. catesbeianus is not well understood although Austin et al. (2004, Mol. Phylogenet. Evol. 32: 799–816) presented mitochondrial DNA evidence of distinct eastern and western lineages. Reviewed by Casper and Hendricks (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 540–546) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 486–515).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. chiricahuensis Chiricahua Leopard Frog

The status of southern Arizona and Mexican populations needs study. Rana subaquavocalis Platz, 1993, is a synonym according to Goldberg et al. (2004, J. Herpetol. 38: 313–319), although some authors (e.g., Hillis and Wilcox, 2005, Mol. Phylogenet. Evol. 34: 299–314; Dubois, 2006, C. R. Biol., Paris 329: 823–840) have continued to recognize the two taxa as distinct species, without comment. Reviewed by Sredl and Jennings (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 546–549, in the sense of including the central Arizona populations now transferred to Lithobates fisheri), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 515–522). See comment under L. fisheri.

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. clamitans Green Frog

Austin and Zamudio (2008, Mol. Phylogenet. Evol. 48: 1041-1053) reported on interpopulational variation at the molecular level and suggested an historical structure inconsistent with the recognized subspecies, which are here rejected on that basis. Reviewed by Stewart (1968, Cat. Am. Amph. Rept. 337), Pauley and Lannoo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 549–552), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 522–547) as Rana clamitans.

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. fisheri Vegas Valley Leopard Frog

Until recently, this species has been considered to be highly restricted in range and extinct. However, Hekkala et al. (2011. Conserv. Genet. 12: 1379–1385) used DNA sequence data from museum specimens to show that L. fisheri and frogs ascribed to R. chiricahuensis from near the Mogollon Rim in central Arizona comprise a lineage that is distinct from R. chiricahuensis populations to the south and east. Platz (1993, J. Herpetol. 27: 154–162) previously noted the various lines of evidence suggesting that L. chiricahuensis was composed of more than one species, with the central Arizona population notably distinctive, but it was not possible, at that time, to compare those frogs genetically with L. fisheri. Reviewed by Jennings (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 554–555, in the sense of only referring to the Vegas Valley population, which is now was then considered to be? extinct) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 547–551).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. grylio Pig Frog

Reviewed by Altig and Lohoefener (1982, Cat. Am. Amph. Rept. 286, as Rana grylio), Richter (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 555–557, as Rana grylio) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 551–556).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. heckscheri River Frog

Reviewed by Sanders (1984, Cat. Am. Amph. Rept. 348) as Rana heckscheri), Butterfield and Lannoo, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 557–558, as Rana heckscheri), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 556–560).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. kauffeldi Mid-Atlantic Coast Leopard Frog

The recognition of this species may require revision of the range of L. pipiens and L. palustris to exclude areas of southern New York, southern Connecticut, Rhode Island, and parts of Massachusetts. The original publication’s association of this species on the basis of molecular data allied this species with Lithobates palustris rather than L. sphenocephalus, suggesting that issues of identification may run deeper than originally suggested.

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-17

l. okaloosae Florida Bog Frog

Reviewed by Moler (1993, Cat. Am. Amph. Rept. 561, as Rana okaloosae) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 561–564). Austin et al. (2003, Biol. J. Linn. Soc. 80: 601–624) discussed the genetic relationship of L. okaloosae and L. clamitans.

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. onca Relict Leopard Frog

The status of this taxon is controversial. Jaeger et al. (2001, Copeia 2001: 339–351) noted a close relationship with L. yavapaiensis, and Pfeiler and Markow (2008, Mol. Phylogenet. Evol. 49: 343-348) reported evidence consistent with a close or identical relationship with L. yavapaiensis. Reviewed by Jennings (1988, Cat. Am. Amph. Rept. 417, as Rana onca) and Bradford, Jennings, and Jaeger (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 567–568) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 565–568).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. palustris Pickerel Frog

Geographic variation studied by Pace (1974, Misc. Publ. Mus. Zool. Univ. Michigan 148). Reviewed by Schaaf and Smith (1971, Cat. Am. Amph. Rept. 117, as Rana palustris) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 568–578).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. pipiens Northern Leopard Frog

Synonymy and discussion in Pace (1974, Misc. Publ. Mus. Zool. Univ. Michigan 148) as Rana pipiens. Reviewed by Rorabaugh (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 570–576) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 578–608).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. septentrionalis Mink Frog

Reviewed by Hedeen (1977, Cat. Am. Amph. Rept. 202, as Rana septentrionalis) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 608–617).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. sevosus Dusky Gopher Frog

Reviewed by Altig and Lohoefener (1983, Cat. Am. Amph. Rept. 324, as Rana areolata sevosa), Richter and Jensen (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 584–586), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 617–621). Recognized as distinct from L. capito and L. areolatus by Young and Crother (2001, Copeia, 2001: 382–388).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. sphenocephalus Southern Leopard Frog

Pace (1974, Misc. Publ. Mus. Zool. Univ. Michigan 148) revived the older name Rana utricularius Harlan, 1825, for this species, which Pace emended to R. utricularia. Subsequently, the International Commission of Zoological Nomenclature moved (Opinion, 1685, 1992, Bull. Zool. Nomencl. 49: 171–173) to suppress R. utricularia for purposes of priority in favor of R. sphenocephala, leaving the unusual situation of the subspecies name sphenocephalus having priority over the older species name, utricularius. The status of the nominal subspecies requires detailed examination (see Brown et al., 1977, Bull. Zool. Nomencl. 33: 199–200; Zug, 1982, Bull. Zool. Nomencl. 39: 80–81; and Uzzell, 1982, Bull. Zool. Nomencl. 39: 83). Reviewed by Butterfield, Lannoo, and Nanjappa (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 586–590) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 621–637). See comment under L. kauffeldi.

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. sphenocephalus sphenocephalus Florida Leopard Frog

Notes on genus: This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. sphenocephalus utricularius Coastal Plains Leopard Frog

Notes on genus: This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. sylvaticus Wood Frog

The extensive morphological variation in this species was examined by Martof and Humphries (1959, Amer. Midl. Nat. 61: 350–389), who rejected previously recognized taxonomic divisions; however a study of DNA sequence variation by Lee-Yaw et al. (2008, Mol. Ecol. 17: 867–884) revealed two distinct clades corresponding to eastern and western populations. Reviewed by Martof (1970, Cat. Am. Amph. Rept. 86, as Rana sylvatica.), Redmer and Trauth (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 590–593, as Rana sylvatica), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 637–669).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. tarahumarae Tarahumara Frog

Extinct in the USA although persisting in Mexico. Attempts are being made to reintroduce the species into former Arizona localities. Reviewed by Zweifel (1968, Cat. Am. Amph. Rept. 66, as Rana tarahumarae.), Rorabaugh and Hale (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 593–595, as Rana tarahumarae), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 669–637).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. virgatipes Carpenter Frog

Reviewed by Gosner and Black (1968, Cat. Am. Amph. Rept. 67, as Rana virgatipes), Mitchell (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 595–596, as Rana virgatipes), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 674–681). Data provided by Pytel (1986, Herpetologica 42: 273–282) suggest that careful evaluation for cryptic species is warranted.

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

l. yavapaiensis Lowland Leopard Frog

See comment under L. onca. Reviewed by Sredl (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 596–599, as Rana yavapaiensis) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 681–636).

Note on genus:
This taxon of North, Central, and South American frogs was removed from the large and predominantly Eurasian genus Rana by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297). Hillis and Wilcox (2005, Mol. Phylogenet. Evol. 34: 299–314) provided a phylogenetic taxonomy that retained the species now under Lithobates within Rana and restricted the use of that name to a small part of what was subsequently assigned to Lithobates by Frost et al. (2006, op. cit.). Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330) criticized the nomenclatural proposals of Hillis and Wilcox and regarded their names as nomina nuda and their approach outside of the International Code of Zoological Nomenclature (1999). This criticism was responded to by Hillis (2006, Mol. Phylogenet. Evol. 42: 331–338), who argued that most of the new names proposed by Hillis and Wilcox do have nomenclatural status under the International Code of Zoological Nomenclature (1999). The revision by Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) which recognized Lithobates as a genus, we think best reflects the majority opinion of members of the international community who are actively working on large-scale ranid relationships, although Hillis, 2007 (Mol. Phylogenet. Evol. 42: 331–338) and Wiens et al. (2009, Evolution 63: 1217–1231) expressed reluctance to accept this taxonomy. Dubois (2006, Mol. Phylogenet. Evol. 42: 317–330; 2007, Cladistics 23: 390–402), Hillis (2007, op. cit.), Pauly et al. (2009, Herpetologica 65: 115–128), Frost et al. (2009, Herpetologica, 65: 136–153) discussed the issues surrounding the nomenclature of North American ranids and most recently Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.: 390–391), suggested that Lithobates be considered a subgenus of Rana. A different approach was suggested by Yuan et al. (Syst. Biol., 65: 824–842) who suggested returning Lithobates to Rana, with Lithobates found to be monophyletic by them, being arrayed as Rana sylvaticus + 4 subgenera within Rana, without applying a name to the overarching Lithobates group. Given that arguments about name stability are largely 10 years and thousands of citations late and also turn on what earlier authors may have meant by “Rana pipiens” when likely few experimental animals with this name attached to them were correctly identified, it seems that the best course of action at this point is to hold this taxonomic change in abeyance.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-01-30

o. septentrionalis Cuban Treefrog

Alien Species:

The Cuban Treefrog is native to Cuba, the Bahamas, and Cayman Islands, has been introduced into eight states, and is established in Florida. It has been claimed to be established in Hawaii (McKeown, 1996, A Field Guide to Reptiles and Amphibians in the Hawaiian Islands, Diamond Head Publishing, Inc., Los Osos, California) but there is no supporting evidence.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

p. brachyphona Mountain Chorus Frog

Reviewed by Hoffmann (1980, Cat. Am. Amph. Rept. 234), Mitchell and Pauley (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 465–466) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 313–318).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. brimleyi Brimley's Chorus Frog

Reviewed by Hoffmann (1983, Cat. Am. Amph. Rept. 311), Mitchell (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 466 - 467) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 319–322).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. cadaverina California Treefrog

Reviewed by Gaudin (1979, Cat. Am. Amph. Rept. 225, as Hyla cadaverina), Ervin (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 467–470) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 322–328). Phillipsen and Metcalf (2009, Mol. Phylogenet. Evol. 53: 152–170) reported on considerable geographic structure at the molecular level among populations.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. clarkii Spotted Chorus Frog

Reviewed by Pierce and Whitehurst (1990, Cat. Am. Amph. Rept. 458), Sredl (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 470–472), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 328–331).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. crucifer Spring Peeper

Moriarty and Cannatella (2004, Mol. Phylogenet. Evol. 30: 409–420) rejected subspecies. Reviewed by Butterfield, Lannoo, and Nanjappa (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 472–474), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 331–348).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. feriarum Upland Chorus Frog

See comment under P. kalmi.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. fouquettei Cajun Chorus Frog

Reviewed by Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 357–363).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. hypochondriaca Baja California Treefrog

Recuero et al. (2006, Mol. Phylogenet. Evol. 39: 293–304) recognized this species as distinct from P. regilla and composed of two subspecies, one of which is extralimital, and whose mutual status is unclear. Barrow et al. (2014, Mol. Phylogenet. Evol. 75: 78-900) suggested that the distinction of P. hypochondriaca and P. sierra, drawn on the basis of mtDNA, was not supported by nuDNA analysis. This suggests that this taxon will ultimately be included in the synonymy of Pseudacris regilla.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. hypochondriaca hypochondriaca Northern Baja California Treefrog

Recuero et al. (2006, Mol. Phylogenet. Evol. 39: 293–304) recognized this species as distinct from P. regilla and composed of two subspecies, one of which is extralimital, and whose mutual status is unclear. Barrow et al. (2014, Mol. Phylogenet. Evol. 75: 78-900) suggested that the distinction of P. hypochondriaca and P. sierra, drawn on the basis of mtDNA, was not supported by nuDNA analysis. This suggests that this taxon will ultimately be included in the synonymy of Pseudacris regilla.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. illinoensis Illinois Chorus Frog

Moriarty and Cannatella (2004, Mol. Phylogenet. Evol. 30: 409–420) discussed the arguable distinctiveness of this taxon with respect to Pseudacris streckeri. Reviewed by Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 363–367).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. kalmi New Jersey Chorus Frog

Platz (1989, Copeia 1989: 704–712) retained P. feriarum and P. kalmi as subspecies of one species but suggested that they might also be distinct species on the basis of data presented by Hedges (1986, Syst. Zool. 35: 1–21). Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) confirmed that P. kalmi and P. feriarum are distinct species although the contact zone between these taxa is poorly understood.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. maculata Boreal Chorus Frog

Considered a species distinct from P. triseriata by Platz (1989, Copeia 1989: 704–712). Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the geographic limits of this species although the evidence based only on mitochondrial DNA variation was not accepted by Green et al. (2014, North American amphibians: distribution and diversity. Univ. of California Press.). Reviewed by Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 371–384).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. nigrita Southern Chorus Frog

Reviewed by Gates (1988, Cat. Am. Amph. Rept. 416), Leja (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 474–475), and Dodd ( 2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 385–390). Subspecies rejected by Moriarty and Cannatella (2004, Mol. Phylogenet. Evol. 30: 409–420).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. ocularis Little Grass Frog

Reviewed by Franz and Chantell (1978, Cat. Am. Amph. Rept. 209, as Limnaoedus ocularis), Jensen, (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 475–477), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 391–395).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. ornata Ornate Chorus Frog

For discussion see Harper (1937, Am. Midl. Nat. 18: 260–272). Reviewed by Jensen (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 477–478), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 395–400).

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. regilla Pacific Treefrog

Recuero et al. (2006, Mol. Phylogenet. Evol. 39: 293–304) redelimited this species and revised its range. Rorabaugh and Lannoo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 478–484) provided a detailed account that summarized the literature (in the sense of including Pseudacris sierra and Pseudacris hypochondriaca. Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 400–416) also provided a review.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. sierra Sierran Treefrog

Recognized as distinct from P. regilla by Recuero et al. (2006, Mol. Phylogenet. Evol. 39: 293–304; 2006, Mol. Phylogenet. Evol. 41: 511). See comment under P. hypochondriaca.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. streckeri Strecker's Chorus Frog

Reviewed by Smith (1966, Cat. Am. Amph. Rept. 27), and Shepard et al. (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 484–485) provided reviews in the sense of including Pseudacris illinoensis. Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 416–421) also provided a review. See comment under Pseudacris illinoensis.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

p. triseriata Western Chorus Frog

See comment under P. maculata. Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the geographic limits of this species based on mitochondrial DNA evidence. Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 421–428) reviewed the species.

Note on genus:
Lemmon et al. (2007, Mol. Phylogenet. Evol. 44: 1068–1082) revised the P. nigrita group (P. brimleyi, P. brachyphona, P. clarkii, P. feriarum, P. kalmi, P. maculata, and P. triseriata and an unnamed species, which was subsequently named as Pseudacris fouquettei). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) deployed a system of subgenera based on the work of Lemmon et al., placing the eastern species in the subgenus Pseudacris and the western members (P. cadaverina, P. hypchondriaca, P. regilla, and P. sierra) in the subgenus Hyliola, and the species P. ocularis and P. crucifer in the subgenus Limnaoedus but we have not adopted subgenera in this list. Duellman et al. (2016, Zootaxa 4104: 1–109) restricted the name Pseudacris to the eastern and Rocky Mountain species related to Pseudacris nigrita and allocated the western species, Pseudacris cadaverina, P. hypochondriaca, P. regilla, and H. sierra to Hyliola Mocquard, 1899. We hold this change in abeyance pending some sense of acceptance within the professional community.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. aurora Northern Red-Legged Frog

Reviewed by Altig and Dumas (1972, Cat. Am. Amph. Rept. 160, in the sense of including R. draytonii), Pearl (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 528–530), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 687–697). Evidence of the distinctiveness of this species from R. draytonii was provided by Hayes and Miyamoto (1984, Copeia 1984: 1018–1022), Shaffer et al. (2004, Mol. Phylogenet. Evol. 13: 2667–2677), Conlon et al. (2006, Peptides 27: 1305–1312), and Pauly et al. (2008, J. Herpetol. 42: 668–679).

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. boylii Foothill Yellow-Legged Frog

Reviewed by Zweifel (1968, Cat. Am. Amph. Rept. 71), Fellers (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 534–536), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 697–707). Molecular study of geographic variation of this rapidly disappearing species should prove illuminating.

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. cascadae Cascades Frog

Reviewed by Altig and Dumas (1971, Cat. Am. Amph. Rept. 105), Pearl and Adams (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 538–540), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 707–715). The disjunct populations should be investigated with respect to call and molecular parameters.

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. draytonii California Red-Legged Frog

See comment under R. aurora. Reviewed by Fellers (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 552–554) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 715–722).

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. luteiventris Columbia Spotted Frog

Green et al. (1996, Evolution 50: 374–390) and Cuellar (1996, Biogeographica 72: 145–150) suggested that R. pretiosa was composed of two sibling species. Subsequently Green et al. (1997, Copeia 1997: 1–8) recognized the eastern and northern form, R. luteiventris, as a species distinct from R. pretiosa. Reviewed by Reaser and Pilliod (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 559–563) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 723–732).

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. muscosa Southern Mountain Yellow-Legged Frog

Reviewed by Zweifel (1968, Cat. Am. Amph. Rept. 65), Vredenburg, Fellers, and Davidson (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 563–566), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 733–739). Vredenburg et al. (2007, J. Zool. 271: 361–374) discussed the systematics of this species and its disappearance from large parts of its former range.

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. pretiosa Oregon Spotted Frog

See comment under R. luteiventris. Reviewed by Pearl and Hayes (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 577–580) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 739–747).

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. sierrae Sierra Nevada Yellow-Legged Frog

Vredenburg et al. (2007, J. Zool. 271: 361–374) recognized this species as distinct from R. muscosa. Reviewed by Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 747–752).

Note on genus:
This large taxon of predominantly Eurasian frogs was redelimited by Frost et al. (2006, Bull. Am. Mus. Nat. Hist., 297, and Che et al. (2007, Mol. Phylogenet. Evol. 42: 1–13) to exclude a number of taxa (e.g., Lithobates, Glandirana, Odorrana, Pelophylax). See Lithobates for most North American species formerly associated with Rana and comments regarding taxonomy.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

r. horribilis Mesoamerican Cane Toad

Recently shown to be a distinct species from R. marina by Acevedo et al. (2016, Zootaxa, 4103: 574–586). Found in South Texas south through Middle America to northwestern Peru. Hero and Stoneham (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 417–422) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 186– 191), provided detailed accounts for the USA under Bufo marinus or Rhinella marina, prior to the partition of the overarching species complex.

Note on genus:
This genus of predominantly South American toads was redelimited by Chaparro et al. (2007, Herpetologica 63: 203–212) to reflect the phylogenetic results of Pramuk (2006, Zool. J. Linn. Soc. 146: 407–452). Van Bocxlaer et al. (2010, Science 327: 679–682) suggested that Rhinella is only distantly related to North American toads of the genera Incilius and Anaxyrus. See comment under Anaxyrus, regarding the treatment of this genus as a subgenus by some.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2014-12-22

r. marina South American Cane Toad

Reviewed (in the sense of including R. horribilis within R. marina) by Easteal (1986, Cat. Am. Amph. Rept. 395, as Bufo marinus). Vallinoto et al. (2010, Zool. Scripta 39: 128–140) suggested on the basis of molecular evidence that the North and Central American population may be a distinct species from the South American populations and subsequently Acevedo et al. (2016, Zootaxa, 4103: 574–586) provided additional morphological and molecular evidence for their distinctiveness, with the largely trans-Andean South American species being the one introduced worldwide, including Hawaii, and the Mesoamerican species extending north into South Texas. The status of introduced Florida populations remains unclear, having been introduced from Colombia where both R. marina and R. horribilis are known.

Note on genus:
This genus of predominantly South American toads was redelimited by Chaparro et al. (2007, Herpetologica 63: 203–212) to reflect the phylogenetic results of Pramuk (2006, Zool. J. Linn. Soc. 146: 407–452). Van Bocxlaer et al. (2010, Science 327: 679–682) suggested that Rhinella is only distantly related to North American toads of the genera Incilius and Anaxyrus. See comment under Anaxyrus, regarding the treatment of this genus as a subgenus by some.

Darrel R. Frost, David M. Green, Emily Moriarty Lemmon, Roy W. McDiarmid, Joseph R. Mendelson III, Fred Kraus, and Kenneth L. Krysko, 2018-02-12

r. dorsalis Burrowing Toad

Geographic variation has not been studied in any detail and cryptic lineages are a possibility. Reviewed by Fouquette (1969, Cat. Am. Amph. Rept. 78) and Fouquette (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 599–600).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. couchii Couch's Spadefoot

Reviewed by Wasserman (1970, Cat. Am. Amph. Rept. 85), Morey (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 508–511), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 753–760). Geographic variation is poorly documented.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. holbrookii Eastern Spadefoot

Reviewed by Wasserman (1968, Cat. Am. Amph. Rept. 70, as Scaphiopus h. holbrookii), Palis (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 511–512), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 772–776).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. hurterii Hurter's Spadefoot

Reviewed by Wasserman (1968, Cat. Am. Amph. Rept. 70, as Scaphiopus holbrookii hurterii), briefly by Lannoo (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 512–513), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 772–776).

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. baudinii Mexican Treefrog

Reviewed by Duellman (1968, Cat. Am. Amph. Rept. 59), Malone (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 489–491), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 431–435).

Note on genus:
The content of this taxon was redelimited by Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist. 294) to include former Pternohyla.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. fodiens Lowland Burrowing Treefrog

Reviewed by Trueb (1969, Cat. Am. Amph. Rept. 77, as Pternohyla fodiens), Sredl (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 488–489), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 428–431).

Note on genus:
The content of this taxon was redelimited by Faivovich et al. (2005, Bull. Am. Mus. Nat. Hist. 294) to include former Pternohyla.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. bombifrons Plains Spadefoot

Known to hybridize with S. multiplicata in parts of their ranges (Brown, 1976, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 286). Geographic variation is poorly documented. Reviewed by Farrar and Hey (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 513–514) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 777–785).

Note on genus:
Tanner (1989, Great Basin Nat. 49: 38–70) and Wiens and Titus (1991, Herpetologica 47: 21–28) recognized Spea as distinct from Scaphiopus, within which it was previously regarded as a subgenus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. hammondii Western Spadefoot

This name formerly covered populations now referred to S. multiplicata and S. intermontana until separated by Brown (1976, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 286). See Tanner (1989, Great Basin Nat. 49: 503–510) for discussion, although he continued to retain these species as subspecies of S. hammondi, a position rejected by Wiens and Titus (1991, Herpetologica 47: 21–38). Reviewed by Morey (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 514–517) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 786–790).

Note on genus:
Tanner (1989, Great Basin Nat. 49: 38–70) and Wiens and Titus (1991, Herpetologica 47: 21–28) recognized Spea as distinct from Scaphiopus, within which it was previously regarded as a subgenus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. intermontana Great Basin Spadefoot

Geographic variation very poorly documented, and, according to evidence provided by Titus and Wiens (1991, Herpetologica 47: 21–29), this nominal species may be a paraphyletic composite of at least two species. Reviewed by Hall (1999, Cat. Am. Amph. Rept. 650), Morey (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 517–519), and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 791–797).

Note on genus:
Tanner (1989, Great Basin Nat. 49: 38–70) and Wiens and Titus (1991, Herpetologica 47: 21–28) recognized Spea as distinct from Scaphiopus, within which it was previously regarded as a subgenus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. multiplicata Mexican Spadefoot

Considered a species distinct from S. hammondii by Brown (1976, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 286) and by Titus and Wiens (1991, Herpetologica 47: 21–28). Regarded, on the basis of overall similarity and paleoclimatic inference to be conspecific with S. hammondii by Van Devender, Mead, and Rea (1991, Southwest. Nat. 36: 302–314) and by Tanner (1989, Great Bas. Nat. 49: 503–510). Tanner recognized S. h. stagnalis Cope as the northern (Arizona to central Chihuahua) subspecies of his Spea hammondii, though the phylogenetic evidence presented by Titus and Wiens indicated it to be part of S. multiplicata. Geographic variation has not been carefully studied and cryptic species are possible. Reviewed by Morey (2005, in Lannoo, M. [ed.], Amphibian Declines: the Conservation Status of United States Species. Univ. of California Press: 519–522) and Dodd (2013, Frogs of the United States and Canada, John Hopkins Univ. Press: 798–806).

Note on genus:
Tanner (1989, Great Basin Nat. 49: 38–70) and Wiens and Titus (1991, Herpetologica 47: 21–28) recognized Spea as distinct from Scaphiopus, within which it was previously regarded as a subgenus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

s. multiplicata stagnalis Chihuahuan Desert Spadefoot

See comment in S. multiplicata.

Note on genus:
Tanner (1989, Great Basin Nat. 49: 38–70) and Wiens and Titus (1991, Herpetologica 47: 21–28) recognized Spea as distinct from Scaphiopus, within which it was previously regarded as a subgenus.

Darrel R. Frost, Emily Moriarty Lemmon, Roy W. McDiarmid, and Joseph R. Mendelson III, 2018-02-06

x. laevis African Clawed Frog

Alien Species:

The African Clawed Frog is native to southern Africa, has been reported from nine states, and is established in Arizona, California, and Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

caudata — salamanders

a. annulatum Ringed Salamander

a. barbouri Streamside Salamander

a. bishopi Reticulated Flatwoods Salamander

Pauly et al. (2006, Mol. Ecol. 16: 415 - 429) recognized western populations of A. cingulatum as a distinct species. They inadvertently reversed the proposed vernacular name with that for A. cingulatum.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. californiense California Tiger Salamander

a. cingulatum Frosted Flatwoods Salamander

Pauly et al. (2006, Mol. Ecol. 16: 415 - 429) recognized western populations of A. cingulatum as a distinct species (A. bishopi) and proposed a new vernacular name for this species. They inadvertently reversed the proposed vernacular name with that for A. bishopi.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. gracile Northwestern Salamander

a. jeffersonianum Jefferson Salamander

Taxonomic recognition of asexual forms that combine genomes of this species, A. laterale, A. texanum, and A. tigrinum raises complex issues. These include discordance between cytoplasmic and nuclear genes, reticulate evolution, and genome-swapping (Bogart, 2003, in Sever, D.M. [ed.], Reproductive Biology and Phylogeny of Urodela, Science Publishers, Inc., Pp. 109-134). Bi and Bogart (2010, BMC Evol. Biol. 10: 238) confirm an ancient origin for the mitochondrial genome shared by asexual forms of this complex. Dubois and Rafaëlli (2012, Alytes 28: 77–161) resurrected the name platineum for the asexual forms.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. laterale Blue-Spotted Salamander

See comment under A. jeffersonianum

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. mabeei Mabee's Salamander

a. macrodactylum Long-Toed Salamander

Lee-Yaw and Irwin (2012, J. Evol. Biol. 25: 2276–2287) and Lee-Yaw et al. (2014, Mol. Ecol. 23: 4590–4602) evaluated geographic variation of mtDNA and nuclear genes throughout the range of the species and found the distributions of five lineages did not completely agree with those of the five presently recognized subspecies, but suggested no changes in the taxonomy of the species.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. macrodactylum columbianum Eastern Long-Toed Salamander

a. macrodactylum croceum Santa Cruz Long-Toed Salamander

a. macrodactylum krausei Northern Long-Toed Salamander

a. macrodactylum macrodactylum Western Long-Toed Salamander

a. macrodactylum sigillatum Southern Long-Toed Salamander

a. maculatum Spotted Salamander

a. mavortium Western Tiger Salamander

Shaffer and McKnight (1996, Evolution 50: 417–433) provided molecular data indicating that the eastern and western tiger salamanders should be regarded as distinct species, and treated the western forms as subspecies of Ambystoma mavortium. Lannoo (2005, in Lannoo M., [ed.], Amphibian Declines, Status of United States Species, Univ. California Press, Pp. 636–639) includes A. mavortium in A. tigrinum.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. mavortium diaboli Gray Tiger Salamander

a. mavortium mavortium Barred Tiger Salamander

a. mavortium melanostictum Blotched Tiger Salamander

a. mavortium nebulosum Arizona Tiger Salamander

a. mavortium stebbinsi Sonoran Tiger Salamander

a. opacum Marbled Salamander

a. talpoideum Mole Salamander

a. texanum Small-Mouthed Salamander

a. tigrinum Eastern Tiger Salamander

See comment under A. mavortium.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. means Two-Toed Amphiuma

a. pholeter One-Toed Amphiuma

a. tridactylum Three-Toed Amphiuma

a. aeneus Green Salamander

a. ferreus Clouded Salamander

a. flavipunctatus Black Salamander

Rissler and Apodaca (2007, Syst. Biol. 56: 924–942) conclude, on the basis of mitochondrial DNA phylogeography and ecological niche modeling, that this taxon should be subdivided into two or more species. Dubois and Raffaëlli (2012, Alytes 28: 77–161) formally recognize A. iecanus Cope, 1883, niger Myers and Maslin, 1948, and a nomen nudum, “sequoiensis” Lowe, 1950, the latter based on an unpublished thesis. Reilly et al. (2013, Diversity 5: 657–679) used nuclear and mitochondrial DNA to show that northern populations in the Klamath, Smith, and Rogue River drainages are genetically distinct from other populations but made no taxonomic changes. Raffaëlli (2013, Les Urodèles du Monde, 2e edition, Penclen Édition, France) recognizes A. niger and A. iëcanus (although diacritical marks are not to be used in scientific names), and also refers to an unnamed subspecies of A. flavipunctatus that he formerly treated as “sequoiensis” by the French vernacular name “Anéides noir de l’Est”. Furthermore, he refers to the populations identified by Reilly et al. (2012, Mol. Ecol. 21: 5745–5761) as an unnamed species, to which he applies the French vernacular name “Anéides noir du Nord”. Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles: The United States and Canada. Xlibris Corporation.) do not recognize the subspecies listed below. Reilly and Wake (2012 Molec. Ecol. 21: 5745–5761), Reilly et al. (2013: Diversity 5: 657–679), and Reilly and Wake (2014, J. Biogeog. 41: 280–2910) suggested that there are undescribed species within A. flavipunctatus on the basis of molecular data, but did not revise the present taxonomy.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

a. flavipunctatus flavipunctatus Speckled Black Salamander

a. flavipunctatus niger Santa Cruz Black Salamander

a. hardii Sacramento Mountains Salamander

a. lugubris Arboreal Salamander

a. vagrans Wandering Salamander

b. altasierrae Greenhorn Mountains Slender Salamander

b. attenuatus California Slender Salamander

This species is highly differentiated with respect to mitochondrial DNA and Martinéz-Solano et al. (2007, Molec. Ecol., 16: 4335 - 4355) recognized five major clades. Highton (2014, Molec. Phylo. and Evol., 71: 127 - 141), using only the mtDNA data, suggested that as many as 39 species should be recognized.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

b. bramei Fairview Slender Salamander

b. campi Inyo Mountains Salamander

b. diabolicus Hell Hollow Slender Salamander

b. gabrieli San Gabriel Mountains Slender Salamander

b. gavilanensis Gabilan Mountains Salamander

b. gregarius Gregarious Slender Salamander

b. incognitus San Simeon Slender Salamander

b. kawia Sequoia Slender Salamander

b. luciae Santa Lucia Mountains Slender Salamander

b. major Southern California Slender Salamander

Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) follow Hansen and Wake (2005, in Lannoo M., [ed.], Amphibian Declines, Status of United States Species, Univ. California Press, Pp. 666–667) in treating this form as a full species. The status of this taxon is considered in Martínez-Solano et al. (2012, Mol. Phylog. Evol. 63: 131–149), who document discordance between nuclear and mitochondrial sequence data in B. major. B. m. aridus is one of six clades of “southern” major with mtDNA data, but “northern” and “southern” components of major are not supported by data from nuclear genes.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

b. major aridus Desert Slender Salamander

Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) follow Hansen and Wake (2005, in Lannoo M., [ed.], Amphibian Declines, Status of United States Species, Univ. California Press, Pp. 666–667) in treating this form as a full species. The status of this taxon is considered in Martínez-Solano et al. (2012, Mol. Phylog. Evol. 63: 131–149), who document discordance between nuclear and mitochondrial sequence data in B. major. B. m. aridus is one of six clades of “southern” major with mtDNA data, but “northern” and “southern” components of major are not supported by data from nuclear genes.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

b. major major Garden Salamander

Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) follow Hansen and Wake (2005, in Lannoo M., [ed.], Amphibian Declines, Status of United States Species, Univ. California Press, Pp. 666–667) in treating this form as a full species. The status of this taxon is considered in Martínez-Solano et al. (2012, Mol. Phylog. Evol. 63: 131–149), who document discordance between nuclear and mitochondrial sequence data in B. major. B. m. aridus is one of six clades of “southern” major with mtDNA data, but “northern” and “southern” components of major are not supported by data from nuclear genes

Stephen G. Tilley (Chair), Richard Highton, David B. Wake, 2014-07-10

b. minor Lesser Slender Salamander

b. nigriventris Black-Bellied Slender Salamander

b. pacificus Channel Islands Slender Salamander

b. regius Kings River Slender Salamander

b. relictus Relictual Slender Salamander

b. robustus Kern Plateau Salamander

b. simatus Kern Canyon Slender Salamander

b. stebbinsi Tehachapi Slender Salamander

b. wrighti Oregon Slender Salamander

c. alleganiensis Hellbender

c. alleganiensis alleganiensis Eastern Hellbender

c. alleganiensis bishopi Ozark Hellbender

Collins (1991, Herpet. Rev. 22: 42–43) elevated this form to species status. Molecular data presented by Crowhurst et al. (2011, Cons. Genet. 12: 637–646) do not support the monophyly of the subspecies, but no formal change in the taxonomy was suggested.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. abditus Cumberland Dusky Salamander

d. aeneus Seepage Salamander

d. apalachicolae Apalachicola Dusky Salamander

d. auriculatus Holbrook's Southern Dusky Salamander

Divergent mitochondrial DNA lineages occur among Atlantic Coastal Plain populations that are morphologically assignable to this species. These lineages do not comprise a monophyletic unit (Beamer and Lamb, 2008, Mol. Phylogenet. Evol. 47: 143–153).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. brimleyorum Ouachita Dusky Salamander

d. carolinensis Carolina Mountain Dusky Salamander

Tilley et al. (2013, Ecol. and Evol., 3: 2547–2567) reported on a molecularly distinctive form in the southern Bald Mountains and northern foothills of the Great Smoky Mountains that is phenotypically indistinguishable from this species. This form appears to hybridize with both D. carolinensis and D. santeetlah in the Blue Ridge Physiographic Province, and with an innominate lowland form further west in the Ridge and Valley Physiographic Province.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. conanti Spotted Dusky Salamander

Populations in the Ridge and Valley Physiographic Province of eastern Tennessee appear to hybridize with this form but Tilley et al. (2013, Ecol. and Evol., 3: 2547–2567) declined to assign them to D. conanti due to their unique mitochondrial haplotypes.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. folkertsi Dwarf Black-Bellied Salamander

d. fuscus Northern Dusky Salamander

Molecular data suggest deep differentiation among populations that morphologically resemble D. fuscus (Bonett, 2002, Copeia 2002: 344–355; Kozak, et al., 2005, Evolution 59: 2000–2016), and additional species almost certainly await resolution.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. imitator Imitator Salamander

d. marmoratus Shovel-Nosed Salamander

Molecular data indicate that this taxon and D. quadramaculatus may not be reciprocally monophyletic (Rissler and Taylor, 2003, Mol. Phylogenet. Evol. 27: 197–211; Kozak, et al., 2005, Evolution 59: 2000–2016; Jones et al. 2006, Mol. Phylogenet. Evol. 38: 280–287; Wooten and Rissler, 2011, Acta Herpetol. 6: 175–208). None of these studies propose taxonomic revisions but Dubois and Rafaëlli (2012, Alytes 28: 77–161) and Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) resurrect two taxa, D.aureatus and D. melanius, from synonymy under this species (Martof , 1962, Am. Midl. Nat., 67: 30).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. monticola Seal Salamander

d. ochrophaeus Allegheny Mountain Dusky Salamander

d. ocoee Ocoee Salamander

This form consists of numerous parapatric units that occupy different mountain ranges in the southern Blue Ridge and Cumberland Plateau physiographic provinces and probably represent distinct species (Tilley and Mahoney, 1996, Herpetol. Monogr. 10: 1–42; Tilley, 1997, J. Heredity 88: 305–315; Highton, 2000, in R. C. Bruce, B. G. Jaeger and L. D, Houck [eds.], The Biology of Plethodontid Salamanders. Kluwer Academic/Plenum Publishers, New York, Pp. 215–241).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. orestes Blue Ridge Dusky Salamander

This taxon consists of two genetically differentiated units that may represent cryptic species (Tilley and Mahoney, 1996, Herpetol. Monogr. 10: 1–42; Tilley, 1997, J. Heredity 88: 305–315; Highton, 2000, in R. C. Bruce, B. G. Jaeger and L. D, Houck [eds.], The Biology of Plethodontid Salamanders. Kluwer Academic/Plenum Publishers, New York, pp. 215–241).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. organi Northern Pygmy Salamander

d. planiceps Flat-Headed Salamander

Removed from synonymy under D. fuscus (Martof and Rose, 1962, Copeia, 1962: 215–216) by Tilley, Eriksen, and Katz (2008, Zool. J. Linnean Soc. 152: 115–130).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

d. quadramaculatus Black-Bellied Salamander

d. santeetlah Santeetlah Dusky Salamander

d. valentinei Valentine's Southern Dusky Salamander

This species was described by Means et al. (2017, Zootaxa 4263: 467-506), on the basis of mtDNA and morphological differences distinguishing it from D. auriculatus.

Richard Highton (Chair), Ronald M. Bonett, Elizabeth L. Jockusch, 2018-01-13

d. welteri Black Mountain Salamander

d. wrighti Pygmy Salamander

d. aterrimus Idaho Giant Salamander

d. copei Cope's Giant Salamander

d. ensatus California Giant Salamander

d. tenebrosus Coastal Giant Salamander

e. eschscholtzii Ensatina

The taxonomy of this complex is controversial. Some authors would recognize from two (e.g., Frost and Hillis, 1990, Herpetologica 46: 87–104) to as many as 11 or more species (e.g., Highton, 1998, Herpetologica 54: 254–278), whereas others (e.g., Wake, 1997, Proc. Natl. Acad. Sci. USA, 94: 7761–7767; Wake and Schneider, 1998, Herpetologica 54: 279–298; Pereira and Wake, 2009, Evolution 68: 2288–2301) consider evidence for evolutionary independence of segments of the complex to be inadequate or equivocal. Narrow hybrid zones have been demonstrated to exist between populations assigned to the subspecies xanthoptica and platensis, and between klauberi and eschscholtzii, and one site of sympatry with no hybridization between the latter pair has been reported (Wake et al., 1989, in D. Otte and J. A. Endler, [eds.], Speciation and its Consequences, Sinauer, Pp. 134–157). Broader zones of genetic admixture and reticulation between units of the complex in many areas raise questions about evolutionary independence, and borders of taxa are elusive.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

e. eschscholtzii croceater Yellow-Blotched Ensatina

e. eschscholtzii eschscholtzii Monterey Ensatina

e. eschscholtzii klauberi Large-Blotched Ensatina

e. eschscholtzii oregonensis Oregon Ensatina

e. eschscholtzii picta Painted Ensatina

e. eschscholtzii platensis Sierra Nevada Ensatina

e. eschscholtzii xanthoptica Yellow-Eyed Ensatina

e. aquatica Brown-Backed Salamander

e. bislineata Northern Two-Lined Salamander

e. chamberlaini Chamberlain's Dwarf Salamander

e. chisholmensis Salado Salamander

e. cirrigera Southern Two-Lined Salamander

e. guttolineata Three-Lined Salamander

e. hillisi Hillis's Dwarf Salamander

This species was described by Wray et al. (2017, Herpetol. Monogr. 31:18-46) based on morphological and mtDNA sequence differences distinguishing it from other Dwarf Salamanders.

Richard Highton (Chair), Ronald M. Bonett, Elizabeth L. Jockusch, 2018-01-13

e. junaluska Junaluska Salamander

e. latitans Cascade Caverns Salamander

e. longicauda Long-Tailed Salamander

e. longicauda longicauda Eastern Long-Tailed Salamander

e. longicauda melanopleura Dark-Sided Salamander

e. lucifuga Cave Salamander

e. multiplicata Many-Ribbed Salamander

Formerly subdivided into the subspecies E. m. griseogaster and E. m. multiplicata. Biochemical data indicate that populations assigned to E. m. griseogaster are conspecific with E. tynerensis, while those of the nominate subspecies fall into two or three divergent clades that may represent distinct species (Bonett and Chippindale, 2004, Mol. Ecol. 13: 1189–1203).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

e. nana San Marcos Salamander

e. naufragia Georgetown Salamander

e. neotenes Texas Salamander

e. paludicola Western Dwarf Salamander

This species was originally described as a subspecies of E. quadridigitata by Mittleman, but he later (1967, Cat. Am. Amphib. Rept 44:1–2) synonymized it with E. quadridigitata. Wray et al. (2017, Herpetol. Monogr.31: 18–46) recognized it as a species on the basis of morphological and mtDNA sequence differences distinguishing it from other Dwarf Salamanders.

Richard Highton (Chair), Ronald M. Bonett, Elizabeth L. Jockusch, 2018-01-13

e. pterophila Fern Bank Salamander

e. quadridigitata Southern Dwarf Salamander

e. rathbuni Texas Blind Salamander

e. robusta Blanco Blind Salamander

e. sosorum Barton Springs Salamander

e. spelaea Grotto Salamander

e. sphagnicola Bog Dwarf Salamander

This species was described by Wray et al. (2017, Herpetol. Monogr. 31:18-46) on the basis of morphological and mtDNA sequence differences distinguishing it from other Dwarf Salamanders.

Richard Highton (Chair), Ronald M. Bonett, Elizabeth L. Jockusch, 2018-01-13

e. subfluvicola Ouachita Streambed Salamander

e. tonkawae Jollyville Plateau Salamander

e. tridentifera Comal Blind Salamander

e. troglodytes Valdina Farms Salamander

e. tynerensis Oklahoma Salamander

e. wallacei Georgia Blind Salamander

This taxon was originally placed in the monotypic genus Haideotriton. It was considered a junior synonym of Eurycea by Dubois (2005, Alytes 23: 20) and shown to nest phylogenetically within Eurycea by Pyron and Wiens (2011, Mol. Phylogenet. Evol. 61: 543–583), and Bonett, et al. (2013 [2014], Evolution 68: 466–482).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

e. waterlooensis Austin Blind Salamander

e. wilderae Blue Ridge Two-Lined Salamander

g. gulolineatus Berry Cave Salamander

Niemiller, et al. (2008, Molec. Ecol. 17: 2258–2275) provide molecular evidence indicating that this form has diverged very recently from G. porphyriticus and is phylogenetically nested within populations referred to that species. Niemiller and Miller (2010, Cat. of American Amph. and Rept. 862: 1–4), Miller and Niemiller (2012, Cat. of American Amph. And Rept. 884: 1–7), and Raffaëlli (2013, Les Urodèles du Monde, 2e edition, Penclen Édition, France – no city given) treat the taxon as a full species, while Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) treat it as a subspecies of G. “porphoriticus”. Bonnet, et al. (2013 [2014], Evolution 68: 466-482) treat it as a subspecies of G. palleucus in their trees but refer to it as a full species in their text. While closely related to G. palleucus, the taxon is distinguished from it in body proportions, osteology, colorations and some genetical aspects (Niemiller and Miller, 2010).

Note on genus: See comment under Pseudotriton montanus. Kuchta et al. (2016, J. Biogeog. 43: 639– 652), in a molecular study of both mitochrondrial and nuclear DNA sequences of the genus, found considerable discordance between their results and the present taxonomy, but did not suggest any taxonomic changes.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

g. palleucus Tennessee Cave Salamander

Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) treat this taxon and its subspecies as subspecies of G. porphyriticus, citing a close relationship to a population of that species suggested in trees in Bonnet, et al. (2013 [2014], Evolution 68: 466-482). That relationship lacks strong statistical support and the latter authors drew no taxonomic conclusions.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

g. palleucus necturoides Big Mouth Cave Salamander

g. palleucus palleucus Pale Salamander

g. porphyriticus Spring Salamander

g. porphyriticus danielsi Blue Ridge Spring Salamander

g. porphyriticus dunni Carolina Spring Salamander

g. porphyriticus duryi Kentucky Spring Salamander

g. porphyriticus porphyriticus Northern Spring Salamander

g. subterraneus West Virginia Salamander

h. scutatum Four-Toed Salamander

Note on genus: Herman and Bouzat (2016, J. Biogeog. 43: 666–678) analyzed geographic variation in a mitochondrial gene throughout the range of the genus and found six highly divergent lineages, but suggested no change in the current taxonomy.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

h. brunus Limestone Salamander

Rovito (2010, Mol. Ecol. 19: 4554–4571) evaluated genetic variation in both mitochondrial and nuclear genes in H. brunus and H. platycephalus and those data supported the hypothesis that H. brunus was derived from H. platycephalus by peripatric speciation. Thus H. platycephalus is paraphyletic, but no changes in its taxonomy were suggested.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

h. platycephalus Mount Lyell Salamander

Rovito (2010, Mol. Ecol. 19: 4554–4571) evaluated genetic variation in both mitochondrial and nuclear genes in H. brunus and H. platycephalus and those data supported the hypothesis that H. brunus was derived from H. platycephalus by peripatric speciation. Thus H. platycephalus is paraphyletic, but no changes in its taxonomy were suggested.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

h. shastae Shasta Salamander

n. alabamensis Black Warrior River Waterdog

Raffaëlli (2013, Les Urodèles du Monde, 2e edition, Penclen Édition, France) and Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris Corporation) recognize N. lodingi, which we treat as a synonym of this form.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

n. beyeri Gulf Coast Waterdog

According to Bart et al. (1997, J. Herpetol. 31: 192–201) this taxon may consist of more than one species.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

n. lewisi Neuse River Waterdog

n. maculosus Mudpuppy

n. maculosus louisianensis Red River Mudpuppy

Collins (1991, Herpet. Rev. 22: 42–43) elevated this form to species rank. Its taxonomic status requires further research.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

n. maculosus maculosus Common Mudpuppy

n. punctatus Dwarf Waterdog

n. meridionalis Black-Spotted Newt

n. meridionalis meridionalis Texas Black-Spotted Newt

n. perstriatus Striped Newt

n. viridescens Eastern Newt

n. viridescens dorsalis Broken-Striped Newt

n. viridescens louisianensis Central Newt

n. viridescens piaropicola Peninsula Newt

n. viridescens viridescens Red-Spotted Newt

p. hubrichti Red Hills Salamander

p. ainsworthi Bay Springs Salamander

This taxon is based on two poorly preserved specimens, one subsequently destroyed, from a single locality in south-central Mississippi. Himes and Beckett (2014, Southeastern Naturalist 12: 851–856) suggest that the taxon be treated as a synonym of Plethodon mississippi, based on their study of the holotype and their inability to find any Plethodon other than P. mississippi at the type locality.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. albagula Western Slimy Salamander

There is molecular and morphological evidence for distinct evolutionary lineages within this taxon (Baird et al., 2006, Copeia 2006: 760–768; Davis and Pauly, 2011, Copeia 2011: 103–112).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. amplus Blue Ridge Gray-Cheeked Salamander

p. angusticlavius Ozark Zigzag Salamander

p. asupak Scott Bar Salamander

p. aureolus Tellico Salamander

p. caddoensis Caddo Mountain Salamander

Shephard and Burbrink (2011, Mol. Phylogenet. Evol. 59: 399–411) sequenced two mitochondrial genes in a detailed study of geographic variation in this species and found four highly divergent groups, but made no taxonomic changes.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. chattahoochee Chattahoochee Slimy Salamander

p. cheoah Cheoah Bald Salamander

p. chlorobryonis Atlantic Coast Slimy Salamander

p. cinereus Eastern Red-Backed Salamander

p. cylindraceus White-Spotted Slimy Salamander

p. dorsalis Northern Zigzag Salamander

p. dunni Dunn's Salamander

p. electromorphus Northern Ravine Salamander

p. elongatus Del Norte Salamander

p. fourchensis Fourche Mountain Salamander

Shephard and Burbrink (2009, Mol. Ecol. 18: 2243–2262) sequenced two mitochondrial genes in a study of geographic variation in the species and found four subgroups but did not suggest changes in the taxonomy.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. glutinosus Northern Slimy Salamander

p. grobmani Southeastern Slimy Salamander

p. hoffmani Valley and Ridge Salamander

p. hubrichti Peaks of Otter Salamander

p. idahoensis Coeur D'alene Salamander

Pelletier et al. (2015, Syst. Biol. 64: 909–925) found support for two independently evolving lineages within this species based on 8 nuclear genes along with mitochondrial data.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. jordani Red-Cheeked Salamander

p. kentucki Cumberland Plateau Salamander

There is molecular evidence for distinct evolutionary lineages within this taxon (Kuchta et al. 2016, PLoS ONE 11(3): 1–25.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. kiamichi Kiamichi Slimy Salamander

p. kisatchie Louisiana Slimy Salamander

p. larselli Larch Mountain Salamander

p. meridianus South Mountain Gray-Cheeked Salamander

p. metcalfi Southern Gray-Cheeked Salamander

p. mississippi Mississippi Slimy Salamander

p. montanus Northern Gray-Cheeked Salamander

p. neomexicanus Jemez Mountains Salamander

p. nettingi Cheat Mountain Salamander

p. ocmulgee Ocmulgee Slimy Salamander

p. ouachitae Rich Mountain Salamander

Shephard and Burbrink (2008, Mol. Ecol. 17: 5315–5335) sequenced two mitochondrial genes in a study of geographic variation in the species and found seven subgroups but did not suggest changes in the taxonomy.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. petraeus Pigeon Mountain Salamander

p. punctatus Cow Knob Salamander

p. richmondi Southern Ravine Salamander

p. savannah Savannah Slimy Salamander

p. sequoyah Sequoyah Slimy Salamander

p. serratus Southern Red-Backed Salamander

Newman and Austin (2015, PLoS ONE: 0130131), and Thesing et al. (2016, Evol. Ecol. 30: 89–104) sequenced mitochondrial DNA from the same or nearby localities in most isolates of this species. They both found five divergent groups, but neither suggested changes in the taxonomy of the group.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. shenandoah Shenandoah Salamander

p. sherando Big Levels Salamander

p. shermani Red-Legged Salamander

p. stormi Siskiyou Mountains Salamander

p. teyahalee Southern Appalachian Salamander

p. vandykei Van Dyke's Salamander

p. variolatus South Carolina Slimy Salamander

p. vehiculum Western Red-Backed Salamander

p. ventralis Southern Zigzag Salamander

p. virginia Shenandoah Mountain Salamander

p. websteri Webster's Salamander

p. wehrlei Wehrle's Salamander

p. welleri Weller's Salamander

p. yonahlossee Yonahlossee Salamander

p. axanthus Southern Dwarf Siren

p. axanthus axanthus Narrow-Striped Dwarf Siren

p. axanthus belli Everglades Dwarf Siren

p. striatus Northern Dwarf Siren

p. striatus lustricolus Gulf Hammock Dwarf Siren

p. striatus spheniscus Slender Dwarf Siren

p. striatus striatus Broad-Striped Dwarf Siren

p. montanus Mud Salamander

Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) transfer this species to the genus Gyrinophilus, citing the cladogram published by Bonnet, et al. (Bonett, et al. (2013 [2014], Evolution 68: 466–482). Those authors, however, refrained from recommending this treatment on the basis of their phylogeny and relationships among forms of Pseudotriton and Gyrinophilus lack strong statistical support in their analysis.

Note on genus: Kozak et al. (2009, Evolution 63: 1769–1784) presented support for the monophyly of Pseudotriton. Bonett et al. (2013 [2014], Evolution 68: 466–482) presented molecular evidence that this genus may not be monophyletic but made no taxonomic recommendations, stating that while Gyrinophilus, Pseudotriton, and Stereochilus form a clade, relationships among these lineages are not well supported in current analyses.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. montanus diastictus Midland Mud Salamander

This taxon was elevated to a full species by Collins (1991, Herpet. Rev. 22: 42–43). This treatment has been followed by Dubois and Raffaëlli (2012, Alytes 28: 77–161) and Raffaëlli (2013, Les Urodèles du Monde, 2e edition, Penclen Édition, France – no city given). Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) also treat it as a subspecies of P. montanus, which they transfer to Gyrinophilus. The phylogeny presented by Bonett, et al. (2013 [2014], Evolution 68: 466–482) indicates a sister relationship between this taxon and P. montanus. In the absence of data on levels of genetic differentiation, we retain the original taxonomic status of this form (Bishop, 1941, Occ. Pap. Mus. Zool. Univ. Mich., 451: 1–27).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. montanus flavissimus Gulf Coast Mud Salamander

Dubois and Raffaëlli (2012, Alytes 28:77–161) consider this taxon a full species, and Raffaëlli (2013, Les Urodèles du Monde, 2e edition, Penclen Édition, France) treats it as a full species, including two subspecies, flavissimus and floridanus.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. montanus floridanus Rusty Mud Salamander

p. montanus montanus Eastern Mud Salamander

p. ruber Red Salamander

Folt et al. (2016, Mol. Phylogen. Evol. 98: 97–110) published an analysis of molecular variation in one nuclear and two mtDNA genes in the southern and western portions of the range of this species. They found several lineages whose geographic ranges do not agree with those of the four presently recognized subspecies. However, they continued to recognize all four subspecies for other reasons.

Note on genus: Kozak et al. (2009, Evolution 63: 1769–1784) presented support for the monophyly of Pseudotriton. Bonett et al. (2013 [2014], Evolution 68: 466–482) presented molecular evidence that this genus may not be monophyletic but made no taxonomic recommendations, stating that while Gyrinophilus, Pseudotriton, and Stereochilus form a clade, relationships among these lineages are not well supported in current analyses.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

p. ruber nitidus Blue Ridge Red Salamander

p. ruber ruber Northern Red Salamander

p. ruber schencki Black-Chinned Red Salamander

p. ruber vioscai Southern Red Salamander

r. cascadae Cascade Torrent Salamander

r. kezeri Columbia Torrent Salamander

r. olympicus Olympic Torrent Salamander

r. variegatus Southern Torrent Salamander

s. intermedia Lesser Siren

S. i. texana was synonymized with S. intermedia nettingi by Flores-Villela and Brandon (1992, Ann. Carnegie Mus. 61: 289–291) but Dubois and Raffaëlli (2012, Alytes 28: 77–161) and Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) considered that subspecies to be valid. The taxonomic statuses of this and the remaining subspecies remain unclear and deserve careful evaluation.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

s. intermedia intermedia Eastern Lesser Siren

s. intermedia nettingi Western Lesser Siren

S. i. texana was synonymized with S. intermedia nettingi by Flores-Villela and Brandon (1992, Ann. Carnegie Mus. 61: 289–291) but Dubois and Raffaëlli (2012, Alytes 28: 77–161) and Fouquette and Dubois (2014, A Checklist of North American Amphibians and Reptiles, The United States and Canada, Seventh Edition, Vol. 1 – Amphibians, Xlibris, San Bernardino, CA, 613 p.) considered that subspecies to be valid. The taxonomic statuses of this and the remaining subspecies remain unclear and deserve careful evaluation.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

s. lacertina Greater Siren

The status of the two distantly allopatric populations (see Flores-Villela and Brandon, 1992, Ann. Carnegie Mus. 61: 289–291) in (1) south Texas and adjacent Mexico and (2) peninsular Florida is unclear and deserves evaluation.

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

s. marginatus Many-Lined Salamander

t. granulosa Rough-Skinned Newt

t. rivularis Red-Bellied Newt

t. sierrae Sierra Newt

Formerly considered a subspecies of T. torosa; elevated to species status by Kuchta (2007, Herpetologica 63: 332–350).

Richard Highton (Chair), Ronald M. Bonnet, Elizabeth L. Jockusch, 2018-02-02

t. torosa California Newt

u. brucei Patch-Nosed Salamander

crocodilia — crocodilians

a. mississippiensis American Alligator

Recent questions concerning the spelling of mississippiensis with “p” or “pp” have led to the inclusion of this annotation with the hope of clearing up the confusion, partly caused by earlier versions of this list. Daudin (1802) spelled it with p and from looking at that volume he probably did it because in French he spelled the river, area, etc., with one p. In 1842 Holbrook (1842, North American Herpetology, or, a Description of the Reptiles inhabiting the United States, 2nd Ed, J. Dobson, Philadelphia) emended the spelling to pp and in 1958 (Hemming 1958, ICZN 1F(F.8):87-126)the ICZN declared the emendation valid. In the meantime, Yarrow (1882, Checklist of North American Batrachia and Reptilia. Bull. U. S. Natl. Mus.) and Cope (1875, Checklist of North American Batrachia and Reptilia. Bull. U. S. Natl. Mus) spelled it pp but Stejneger and Barbour (1917, 1923, 1933, Editions 1-3, A check list of North American amphibians and reptiles. Harvard University Press, Cambridge, Massachusetts), Schmidt (1953, A check list of North American amphibians and reptiles. Sixth Edition. ASIH, Chicago) and Conant et al. (1956, Copeia 1956: 172-185) spelled it with one p. The Collins' lists (1978, 1982, 1990, 1997, Editions 1-4, Standard common and current scientific names for North American amphibians and reptiles. SSAR Herpetological Circular) spelled it with pp but the 5th and 6th editions (Crother et al., 2000, 2008, Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding. SSAR Herpetological Circular 29, 37) went back to p (in error). The spelling was corrected to pp in the most recent (7th) edition (Crother et al., 2012, Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding. SSAR Herpetological Circular 39).

Brian I. Crother, 12-12-14

c. crocodilus Spectacled Caiman

Alien Species:

The Spectacled Caiman is native to South America, has been reported from seven states, and is established in Florida.

Fred Kraus, 2015-01-19

c. acutus American Crocodile

serpentes — snake

a. javanicus Javanese File Snake

Alien Species:

The Javanese File Snake is native to Southeast Asia and is established in Florida.

Fred Kraus, 2015-01-19

a. conanti Florida Cottonmouth

Using multi-locus nuclear data, Burbrink and Guiher (2015, Zool. J. Linn. Soc. 173:505– 526) partially confirmed previous mitochondrial hypotheses (Guiher and Burbrink 2008, Mol. Phylogenet. Evol. 48: 112–125) that the two North American species (A. contortrix and A. piscivorus) each consist of multiple species-level taxa. Subspecies are not recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

a. contortrix Eastern Copperhead

Using multi-locus nuclear data, Burbrink and Guiher (2015, Zool. J. Linn. Soc. 173:505– 526) partially confirmed previous mitochondrial hypotheses (Guiher and Burbrink 2008, Mol. Phylogenet. Evol. 48: 112–125) that the two North American species (A. contortrix and A. piscivorus) each consist of multiple species-level taxa. Subspecies are not recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

a. laticinctus Broad-Banded Copperhead

Using multi-locus nuclear data, Burbrink and Guiher (2015, Zool. J. Linn. Soc. 173:505– 526) partially confirmed previous mitochondrial hypotheses (Guiher and Burbrink 2008, Mol. Phylogenet. Evol. 48: 112–125) that the two North American species (A. contortrix and A. piscivorus) each consist of multiple species-level taxa. Subspecies are not recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

a. piscivorus Northern Cottonmouth

Using multi-locus nuclear data, Burbrink and Guiher (2015, Zool. J. Linn. Soc. 173:505– 526) partially confirmed previous mitochondrial hypotheses (Guiher and Burbrink 2008, Mol. Phylogenet. Evol. 48: 112–125) that the two North American species (A. contortrix and A. piscivorus) each consist of multiple species-level taxa. Subspecies are not recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

a. elegans Glossy Snake

Collins (1991, Herpetol. Rev. 22: 42–43) elevated A. e. occidentalis to specific statusto include all populations in the Sonoran and Mohave Desert regions, the first use of this binomial. Liner (1994, SSAR Herpetol. Circ. 23: 1–113) and Collins (1997, SSAR Herpetol. Circ. 25: 1–40) followed this arrangement. Because no discussion of the taxonomic diagnosis was presented (although Dixon [1959, Southwest. Nat. 4: 20–29] found tail length differences between eastern and western groups), we retain occidentalis as a nominal subspecies.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

a. elegans arenicola Texas Glossy Snake

a. elegans candida Mohave Glossy Snake

a. elegans eburnata Desert Glossy Snake

a. elegans elegans Kansas Glossy Snake

a. elegans noctivaga Arizona Glossy Snake

a. elegans occidentalis California Glossy Snake

Collins (1991, Herpetol. Rev. 22: 42–43) elevated A. e. occidentalis to specific status to include all populations in the Sonoran and Mohave Desert regions, the first use of this binomial. Liner (1994, SSAR Herpetol. Circ. 23: 1–113) and Collins (1997, SSAR Herpetol. Circ. 25: 1–40) followed this arrangement. Because no discussion of the taxonomic diagnosis was presented (although Dixon [1959, Southwest. Nat. 4: 20–29] found tail length differences between eastern and western groups), we retain occidentalis as a nominal subspecies.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

a. elegans philipi Painted Desert Glossy Snake

b. constrictor Boa Constrictor

Alien Species:

The Boa Constrictor is native to Central and South America, has been reported from 12 states, and is established in Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-14

b. rosaliae Baja California Ratsnake

Recognition of Bogertophis as distinct from Elaphe has been corroborated by multiple studies using nuclear and mitochondrial data (Utiger et al., 2002, Russian J. Herpetol. 9: 105–124; Burbrink and Lawson, 2007, Mol. Phylogenet. Evolution 43: 173–189; Pyron and Burbrink, 2009, Mol. Phylogenet. Evol. 52: 524–529).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

b. subocularis Trans-Pecos Ratsnake

Recognition of Bogertophis as distinct from Elaphe has been corroborated by multiple studies using nuclear and mitochondrial data (Utiger et al., 2002, Russian J. Herpetol. 9: 105–124; Burbrink and Lawson, 2007, Mol. Phylogenet. Evolution 43: 173–189; Pyron and Burbrink, 2009, Mol. Phylogenet. Evol. 52: 524–529).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

b. subocularis subocularis Northern Trans-Pecos Ratsnake

Recognition of Bogertophis as distinct from Elaphe has been corroborated by multiple studies using nuclear and mitochondrial data (Utiger et al., 2002, Russian J. Herpetol. 9: 105–124; Burbrink and Lawson, 2007, Mol. Phylogenet. Evolution 43: 173–189; Pyron and Burbrink, 2009, Mol. Phylogenet. Evol. 52: 524–529).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. amoenus Common Wormsnake

c. amoenus amoenus Eastern Wormsnake

c. amoenus helenae Midwestern Wormsnake

c. vermis Western Wormsnake

Clark (1968, Herpetologica 24: 104–112) recommended elevating C. (a.) vermis to species status on the basis of allopatry and morphological differences, but Rossman (1973, J. Herpetol. 7: 140–141) presented evidence for the conspecificity of amoenus and vermis in the form of intergrade populations. Collins (1991, Herpetol. Rev. 22: 42–43) considered C. vermis to be distinct from C. amoenus, implying that the populations discussed by Rossman were either/part of C. vermis, or an unnamed taxon. We follow Clark (1968) but anticipate results from molecular studies to better understand population structure and gene flow among allopatric lineages.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. coccinea Scarletsnake

Last reviewed by Williams and Wilson (1967, Tulane Studies in Zoology 13: 103–124).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. coccinea coccinea Florida Scarletsnake

Last reviewed by Williams and Wilson (1967, Tulane Studies in Zoology 13: 103–124).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. coccinea copei Northern Scarletsnake

Last reviewed by Williams and Wilson (1967, Tulane Studies in Zoology 13: 103–124).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. lineri Texas Scarletsnake

Weinell and Austin (2017. J. Herpetol. 51: 161–171.) elevated lineri based on DNA, color pattern, and lepidosis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-17

c. bottae Northern Rubber Boa

Kluge (1993, Zool. J. Linn. Soc. 107: 293–351) placed Lichanura in the synonymy of Charina because they formed sister taxa. Burbrink (2005, Mol. Phylogenet. Evo. 34: 167–180) corroborated the relationship found by Kluge. Rodríguez-Robles et al. (2001, Mol. Phylogenet. Evol. 18: 227–237) found C. b. umbratica to be morphologically and geographically distinct and were elevated to species status based in part on lineages using mtDNA evidence along with with allozyme data from a previous study (Weisman, 1988, MS Thesis, CSU Polytechnic Pomona). With the recognition of C. umbratica and fossil species referred to both Charina and Lichanura (Holman, 2000, Fossil Snakes of North America, Indiana Univ. Press), neither genus is monotypic, and they are treated here as separate genera.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. umbratica Southern Rubber Boa

Kluge (1993, Zool. J. Linn. Soc. 107: 293–351) placed Lichanura in the synonymy of Charina because they formed sister taxa. Burbrink (2005, Mol. Phylogenet. Evo. 34: 167–180) corroborated the relationship found by Kluge. Rodríguez-Robles et al. (2001, Mol. Phylogenet. Evol. 18: 227–237) found C. b. umbratica to be morphologically and geographically distinct and were elevated to species status based in part on lineages using mtDNA evidence along with with allozyme data from a previous study (Weisman, 1988, MS Thesis, CSU Polytechnic Pomona). With the recognition of C. umbratica and fossil species referred to both Charina and Lichanura (Holman, 2000, Fossil Snakes of North America, Indiana Univ. Press), neither genus is monotypic, and they are treated here as separate genera.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. stramineus Variable Sandsnake

Grismer et al. (2002, Herpetologica 58: 18–31) found the previously recognized species C. cinctus, C. punctatissimus, and C. stramineus to represent morphotypes of a single species.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. annulata Resplendent Desert Shovel-Nosed Snake

There is some question as to the validity of the name C. saxatilis (Funk, 1967, Southwest Nat. 12: 180), the Gila Mountains Shovel-nosed Snake, which is generally considered to be a synonym of C. o. annulata (see Cross, 1978, Ph.D. dissertation, Univ. Arizona). Mahrdt et al. (2001, Cat. Am. Amph. Rept. 730) considered C. saxatilis a synonym of C. o. annulata. Wood et al. (2008, Conserv. Gen. 9: 1489–1507) demonstrated, using mtDNA and morphological data, that population structure was not concordant with the traditional subspecific taxonomy. They also revealed two potentially independent evolutionary lineages. A phylogeographic study from Wood et al. (2014; PLoS ONE e97494) using mtDNA and microsatellites indicates that C. o. annulata should be elevated to species status, while retaining two subspecies C. a. annulata and C. a. klauberi, that conform to patterns of genetic structure. The authors find no support for C. o. talpina and place it in synonomy with C. occipitalis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

c. annulata annulata Colorado Desert Shovel-Nosed Snake

There is some question as to the validity of the name C. saxatilis (Funk, 1967, Southwest Nat. 12: 180), the Gila Mountains Shovel-nosed Snake, which is generally considered to be a synonym of C. o. annulata (see Cross, 1978, Ph.D. dissertation, Univ. Arizona). Mahrdt et al. (2001, Cat. Am. Amph. Rept. 730) considered C. saxatilis a synonym of C. o. annulata. Wood et al. (2008, Conserv. Gen. 9: 1489–1507) demonstrated, using mtDNA and morphological data, that population structure was not concordant with the traditional subspecific taxonomy. They also revealed two potentially independent evolutionary lineages. A phylogeographic study from Wood et al. (2014; PLoS ONE e97494) using mtDNA and microsatellites indicates that C. o. annulata should be elevated to species status, while retaining two subspecies C. a. annulata and C. a. klauberi, that conform to patterns of genetic structure. The authors find no support for C. o. talpina and place it in synonomy with C. occipitalis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

c. annulata klauberi Tucson Shovel-Nosed Snake

c. occipitalis Mohave Shovel-Nosed Snake

There is some question as to the validity of the name C. saxatilis (Funk, 1967, Southwest Nat. 12: 180), the Gila Mountains Shovel-nosed Snake, which is generally considered to be a synonym of C. o. annulata (see Cross, 1978, Ph.D. dissertation, Univ. Arizona). Mahrdt et al. (2001, Cat. Am. Amph. Rept. 730) considered C. saxatilis a synonym of C. o. annulata. Wood et al. (2008, Conserv. Gen. 9: 1489–1507) demonstrated, using mtDNA and morphological data, that population structure was not concordant with the traditional subspecific taxonomy. They also revealed two potentially independent evolutionary lineages. A phylogeographic study from Wood et al. (2014; PLoS ONE e97494) using mtDNA and microsatellites indicates that C. o. annulata should be elevated to species status, while retaining two subspecies C. a. annulata and C. a. klauberi, that conform to patterns of genetic structure. The authors find no support for C. o. talpina and place it in synonomy with C. occipitalis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. palarostris Sonoran Shovel-Nosed Snake

c. palarostris organica Organ Pipe Shovel-Nosed Snake

c. kirtlandii Kirtland's Snake

c. bilineatus Sonoran Whipsnake

Contrary to Collins (1997, SSAR Herpetol. Circ. 25: 1–40), Camper and Dixon (1994, Ann. Carnegie Mus. Nat. Hist. 63: 1–48) did not recognize any subspecies for bilineatus.

Note on genus: Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the New World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13: 93), though based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. constrictor North American Racer

Fitch et al. (1981, Trans, Kansas Acad. Sci. 84: 196–203) argued for the elevation of C. c. mormon. This recommendation was rejected by Greene (1983, J. Herpetol. 18: 210–211), and was supported by Corn and Bury (1986, Herpetologica 42: 258–264), who showed a broad zone of intergradation across Colorado and Utah. Collins (1991, Herpetol. Rev. 22: 42–43) re-elevated mormon to specific status, although allopatry was not suitably demonstrated. Anderson (1996, MS thesis, Southeastern Louisiana Univ.) argued that based on allozyme data C. c. mormon cannot be differentiated but that C. c. paludicola and C. c. oaxaca were diagnosable and should be elevated to species status. We retain C. c. mormon and await action on oaxaca and paludicola until the data are published. Burbrink et al. (2008, Mol. Phylogen. Evol 47: 274–288) have demonstrated using mtDNA that C. constrictor may be composed of six independently evolving lineages not concordant with most recognized subspecies. In particular, neither C. c. mormon or C. paludicola represents an evolutionarily distinct lineage. No samples of C. c. oaxaca were included.

Note on genus: Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the New World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13: 93), though based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. constrictor anthicus Buttermilk Racer

c. constrictor constrictor Northern Black Racer

c. constrictor etheridgei Tan Racer

c. constrictor flaviventris Eastern Yellow-Bellied Racer

c. constrictor foxii Blue Racer

c. constrictor helvigularis Brown-Chinned Racer

c. constrictor latrunculus Black-Masked Racer

c. constrictor mormon Western Yellow-Bellied Racer

Fitch et al. (1981, Trans, Kansas Acad. Sci. 84: 196–203) argued for the elevation of C. c. mormon. This recommendation was rejected by Greene (1983, J. Herpetol. 18: 210–211), and was supported by Corn and Bury (1986, Herpetologica 42: 258–264), who showed a broad zone of intergradation across Colorado and Utah. Collins (1991, Herpetol. Rev. 22: 42–43) re-elevated mormon to specific status, although allopatry was not suitably demonstrated. Anderson (1996, MS thesis, Southeastern Louisiana Univ.) argued that based on allozyme data C. c. mormon cannot be differentiated but that C. c. paludicola and C. c. oaxaca were diagnosable and should be elevated to species status. We retain C. c. mormon and await action on oaxaca and paludicola until the data are published. Burbrink et al. (2008, Mol. Phylogen. Evol 47: 274–288) have demonstrated using mtDNA that C. constrictor may be composed of six independently evolving lineages not concordant with most recognized subspecies. In particular, neither C. c. mormon or C. paludicola represents an evolutionarily distinct lineage. No samples of C. c. oaxaca were included.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. constrictor oaxaca Mexican Racer

Fitch et al. (1981, Trans, Kansas Acad. Sci. 84: 196–203) argued for the elevation of C. c. mormon. This recommendation was rejected by Greene (1983, J. Herpetol. 18: 210–211), and was supported by Corn and Bury (1986, Herpetologica 42: 258–264), who showed a broad zone of intergradation across Colorado and Utah. Collins (1991, Herpetol. Rev. 22: 42–43) re-elevated mormon to specific status, although allopatry was not suitably demonstrated. Anderson (1996, MS thesis, Southeastern Louisiana Univ.) argued that based on allozyme data C. c. mormon cannot be differentiated but that C. c. paludicola and C. c. oaxaca were diagnosable and should be elevated to species status. We retain C. c. mormon and await action on oaxaca and paludicola until the data are published. Burbrink et al. (2008, Mol. Phylogen. Evol 47: 274–288) have demonstrated using mtDNA that C. constrictor may be composed of six independently evolving lineages not concordant with most recognized subspecies. In particular, neither C. c. mormon or C. paludicola represents an evolutionarily distinct lineage. No samples of C. c. oaxaca were included.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. constrictor paludicola Everglades Racer

Fitch et al. (1981, Trans, Kansas Acad. Sci. 84: 196–203) argued for the elevation of C. c. mormon. This recommendation was rejected by Greene (1983, J. Herpetol. 18: 210–211), and was supported by Corn and Bury (1986, Herpetologica 42: 258–264), who showed a broad zone of intergradation across Colorado and Utah. Collins (1991, Herpetol. Rev. 22: 42–43) re-elevated mormon to specific status, although allopatry was not suitably demonstrated. Anderson (1996, MS thesis, Southeastern Louisiana Univ.) argued that based on allozyme data C. c. mormon cannot be differentiated but that C. c. paludicola and C. c. oaxaca were diagnosable and should be elevated to species status. We retain C. c. mormon and await action on oaxaca and paludicola until the data are published. Burbrink et al. (2008, Mol. Phylogen. Evol 47: 274–288) have demonstrated using mtDNA that C. constrictor may be composed of six independently evolving lineages not concordant with most recognized subspecies. In particular, neither C. c. mormon or C. paludicola represents an evolutionarily distinct lineage. No samples of C. c. oaxaca were included.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. constrictor priapus Southern Black Racer

c. flagellum Coachwhip

Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the new World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13:93), albeit based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. flagellum cingulum Sonoran Coachwhip

c. flagellum flagellum Eastern Coachwhip

c. flagellum lineatulus Lined Coachwhip

c. flagellum piceus Red Racer

c. flagellum ruddocki San Joaquin Coachwhip

c. flagellum testaceus Western Coachwhip

c. fuliginosus Baja California Coachwhip

On the basis of a sympatric occurrence with C. flagellum, Grismer (1994, Herpetol. Nat. Hist. 2: 51; 2002, Amphibians and Reptiles of Baja California, Including Its Pacific Islands and the Islands in the Sea of Cortés, Univ. California Press) elevated C. f. fuliginosus to species status.

Note on genus: Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the new World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13:93), albeit based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. lateralis Striped Racer

Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the new World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13:93), albeit based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. lateralis euryxanthus Alameda Striped Racer

c. lateralis lateralis California Striped Racer

c. schotti Schott's Whipsnake

Camper and Dixon (1994, Ann. Carnegie Mus. Nat. Hist. 63: 1–48) elevated C.schotti from C. taeniatus with ruthveni retained as a subspecies.

Note on genus: Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the new World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13:93), albeit based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. schotti ruthveni Ruthven's Whipsnake

Camper and Dixon (1994, Ann. Carnegie Mus. Nat. Hist. 63: 1–48) elevated C.schotti from C. taeniatus with ruthveni retained as a subspecies.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. schotti schotti Schott's Striped Whipsnake

c. taeniatus Striped Whipsnake

Camper and Dixon (1994, Ann. Carnegie Mus. Nat. Hist. 63: 1–48) elevated C.schotti from C. taeniatus with ruthveni retained as a subspecies.

Note on genus: Nagy et al. (2004, J. Zool. Syst. Evol. Res. 42: 223–233) restricted the genus Coluber to the new World and suggested that Masticophis might be paraphyletic with respect Coluber. Utiger et al. (2005, Russian J. Herpetol. 12: 39–60) corroborated Nagy et al., finding Masticophis to be paraphyletic with respect to Coluber and synonymizing Masticophis with Coluber (the oldest available name). This arrangement was also recovered in a recent phylogeny of Squamata (Pyron et al., 2013 BMC Evol. Biol. 13:93), albeit based on much of the same data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. taeniatus girardi Central Texas Whipsnake

c. taeniatus taeniatus Desert Striped Whipsnake

c. imperialis Regal Black-Striped Snake

c. imperialis imperialis Tamaulipan Black-Striped Snake

c. longicauda Forest Sharp-Tailed Snake

This species was originally named Contia longicaudae by Feldman and Hoyer (2010, Copeia, 2010: 254–267); however, because they explicitly treated the second part of the binomen as an adjective, it must agree with the name Contia in gender and number so that the correct spelling is Contia longicauda.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. tenuis Common Sharp-Tailed Snake

c. adamanteus Eastern Diamond-Backed Rattlesnake

Notes on the Genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. atrox Western Diamond-Backed Rattlesnake

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. cerastes Sidewinder

Douglas et al. (2006, Mol. Ecol. 15: 3353–3374), using mtDNA, found several geographically distinct lineages within C. cerastes. Only one of these lineages corresponded to a recognized subspecies. (C. c. laterorepens).

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. cerastes cerastes Mohave Desert Sidewinder

c. cerastes cercobombus Sonoran Sidewinder

c. cerastes laterorepens Colorado Desert Sidewinder

Douglas et al. (2006, Mol. Ecol. 15: 3353–3374), using mtDNA, found several geographically distinct lineages within C. cerastes. Only one of these lineages corresponded to a recognized subspecies. (C. c. laterorepens).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. cerberus Arizona Black Rattlesnake

See annotation under C. oreganus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. horridus Timber Rattlesnake

Pisani et al. (1972, Trans. Kansas Acad. Sci. 75: 255–263) conducted a multivariate analysis of variation in C. horridus and concluded that characters tended to be clinal and recommended against recognition of the two subspecies. Brown and Ernst (1986, Brimleyana 12: 57–74) countered that morphology in the eastern part of the range supported recognition of coastal plain and montane subspecies. Clark et al. (2003, J. Herpetol. 37: 145–154) identified three mitochondrial DNA lineages separated by the Appalachian and Allegheny Mountain ranges that did not correspond with the classic arrangement of subspecies within C. horridus.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. lepidus Rock Rattlesnake

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. lepidus klauberi Banded Rock Rattlesnake

c. lepidus lepidus Mottled Rock Rattlesnake

c. molossus Western Black-Tailed Rattlesnake

The northern populations of this species were examined in detail using a multi-locus nuclear dataset (Anderson and Greenbaum, 2013; Herp. Monogr. 26: 19–57), supporting recognition of C. molossus for populations west of the Cochise Filter Barrier (from the Sonoran Desert west), and C. ornatus for eastern populations (from Chihuahuan Desert east), with a narrow contact zone.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. oreganus Western Rattlesnake

Pook et al. (2000, Mol. Phylogenet. Evol. 15: 269–282), Ashton and de Queiroz (2001, Mol. Phylogenet. Evol. 21: 176–189), and Douglas et al. (2002, Biology of the Vipers, Schuett, Hoggren, Douglas, Greene [eds.] Eagle Mountain Press) analyzed mtDNA sequence data and concluded that Crotalus viridis comprised at least two clades, C. viridis and C. oreganus, with C. cerberus being the sister taxon to populations of C. oreganus. The former two studies did not formally recognize C. cerberus as a species, although both suggested that it was distinct based on sequence differences and allopatry. The latter study did recognize C. cerberus as well as four other taxa. Although the studies relied on the same locus, we conservatively conclude that the congruence among all three studies might suggest the recognition of C. viridis, C. oreganus and C. cerberus. A recent unpublished study (Goldenberg, 2013; MS Dissertation, SDSU, 90 pp.) suggests a unique lineage, that has not yet been named, occurs in the southern part of the nominate species' range, and that the subspecies as currently recognized do not correspond with the actual species-level divergences in the group.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. oreganus abyssus Grand Canyon Rattlesnake

c. oreganus concolor Midget Faded Rattlesnake

c. oreganus helleri Southern Pacific Rattlesnake

c. oreganus lutosus Great Basin Rattlesnake

c. oreganus oreganus Northern Pacific Rattlesnake

c. ornatus Eastern Black-Tailed Rattlesnake

The northern populations of this species were examined in detail using a multi-locus nuclear dataset (Anderson and Greenbaum, 2013; Herp. Monogr. 26: 19–57), supporting recognition of C. molossus for populations west of the Cochise Filter Barrier (from the Sonoran Desert west), and C. ornatus for eastern populations (from Chihuahuan Desert east), with a narrow contact zone.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. pricei Twin-Spotted Rattlesnake

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. pricei pricei Western Twin-Spotted Rattlesnake

c. pyrrhus Southwestern Speckled Rattlesnake

Meik et al. (2015. PLoS ONE 10(6): e0131435. doi: 10.1371/journal.pone.0131435), using multilocus sequence and phenotypic data, demonstrated that C. pyrrhus is a species distinct from C. mitchellii of Baja California.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

c. ruber Red Diamond Rattlesnake

The International Commission on Zoological Nomenclature (2000, Bull. Zool. Nomencl. 57: 189–190. Opinion 1960) has ruled that the name Crotalus ruber Cope 1892 take precedence over C. exsul Garman 1884 when used as a specific epithet.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. scutulatus Mohave Rattlesnake

The spelling of the word “Mojave” or “Mohave” has been a subject of debate. Lowe in the preface to his “Venomous Reptiles of Arizona” (1986) argued for “Mohave” as did Campbell and Lamar (2004, The Venomous Reptiles of the Western Hemisphere, Comstock Publishing). According to linguistic experts on Native American languages, either spelling is correct, but using either the “j” or “h” is based on whether the word is used in a Spanish or English context. Given that this is an English names list, we use the “h” spelling (P. Munro, Linguistics, UCLA, pers. comm.). Jones (2016, Sonoran Herpetol. 29: 64–71) argued that the spelling should be with “j” but the committee was not convinced and voted to continue to spell it as Mohave.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. scutulatus scutulatus Northern Mohave Rattlesnake

The English name of the nominal subspecies has been changed to reflect the distribution rather than describe rattlesnakes from a small portion of its distribution (D. Hardy and H. Greene, pers. comm.)

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

c. stephensi Panamint Rattlesnake

Elevated to species by Douglas et al. (2007, Copeia 2007 (4): 920–932).

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. tigris Tiger Rattlesnake

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. viridis Prairie Rattlesnake

See comments under C. oreganus. Douglas et al. (2002, Biology of the Vipers, Schuett, Hoggren, Douglas, Greene [eds.] Eagle Mountain Press) synonymized C. v. nuntius with C. v. viridis.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. willardi Ridge-Nosed Rattlesnake

Barker (2016, in Schuett et al. [eds] Rattlesnakes of Arizona. ECO Publishing) recommended elevating the five subspecies of C. willardi to species, which we do not follow until data are available for evaluation.

Notes on genus: The traditional view of rattlesnake taxonomy that recognizes the two monophyletic sister genera Crotalus and Sistrurus (e.g. Brattstrom, 1964, San Diego Soc. Nat. Hist. 13: 185–268) has recently been challenged. Stille (1987, Herpetologica 43: 98–104) and McCranie (1989, Herpetologica 44: 123–126) presented data that suggested Sistrurus is not monophyletic and rendered Crotalus paraphyletic. Parkinson (1999, Copeia 1999: 576–586) found Sistrurus monophyletic but its position rendered Crotalus paraphyletic. Knight et al. (1993, Syst. Biol. 42: 356–367) used mtDNA to defend the traditional generic taxonomy, but in order to do so ignored the most parsimonious tree. The genus Crotalus is monophyletic when including the Mexican C. ravus (Murphy et al. 2002, in Schuett et al. [eds.] Biology of the Vipers, Eagle Mountain Publishing, Pp. 69–92), and is supported as such in most recent phylogenies, as well as being the sister taxon to a monophyletic Sistrurus (e.g., Pyron et al., 2013; BMC Evol. Biol. 13: 93). Davis et al. (2016. PLoS ONE 11(1): e0146166.doi:10.1371/journal.pone.0146166) used mtDNA and morphometric analyses that resolved six species within the C. viridis complex, which we don’t follow pending further analyses with nDNA.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

c. willardi obscurus New Mexico Ridge-Nosed Rattlesnake

c. willardi willardi Arizona Ridge-Nosed Rattlesnake

d. punctatus Ring-Necked Snake

Numerous data suggest that more than one lineage exists (Blanchard, 1942, Bull. Chicago Acad. Sci. 7: 1–144; Gelbach, 1974, Herpetologica 30: 140–148; Pinou et al., 1995, J. Herpetol. 29: 105–110; Feldman and Spicer, 2006, Mol. Ecol. 15: 2201–2222). Using mitochondrial data sampled from specimens across their range, Fontanella et al. (2008, Mol. Phylogenet. Evol. 46: 1049–1070) found at least 14 lineages that do not follow the geographic range of the subspecies, and may be independently evolving taxa. While D. punctatus may be divided into several species in the near future, we refrain from making any changes at present. Evidence to synonymize the various races into a single species has been poorly presented, and our arrangement follows the traditional subspecies groupings.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

d. punctatus acricus Key Ring-Necked Snake

d. punctatus amabilis Pacific Ring-Necked Snake

d. punctatus arnyi Prairie Ring-Necked Snake

d. punctatus edwardsii Northern Ring-Necked Snake

d. punctatus modestus San Bernardino Ring-Necked Snake

d. punctatus occidentalis Northwestern Ring-Necked Snake

d. punctatus pulchellus Coral-bellied Ring-necked Snake

d. punctatus punctatus Southern Ring-Necked Snake

d. punctatus regalis Regal Ring-Necked Snake

d. punctatus similis San Diego Ring-Necked Snake

d. punctatus stictogenys Mississippi Ring-Necked Snake

d. punctatus vandenburgii Monterey Ring-Necked Snake

d. couperi Eastern Indigo Snake

Wüster et al. (2001, Herpetol. J. 11: 157–165) demonstrated that couperi is a distinct species using morphological evidence.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

d. kolpobasileus Gulf Coast Indigo Snake

d. melanurus Central American Indigo Snake

Wüster et al. (2001, Herpetol. J. 11: 157–165) showed that the South American D. corais is distinct from the Central/North American (D. melanurus) taxon.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

d. melanurus erebennus Texas Indigo Snake

d. margaritiferus Speckled Racer

d. margaritiferus margaritiferus Northern Speckled Racer

f. abacura Red-Bellied Mudsnake

Cundall and Rossman (1984, Herpetologica 40: 388–405) analyzed skull morphology and showed substantial divergence between F. a. abacura and F. a. reinwardtii.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

f. abacura abacura Eastern Mudsnake

Cundall and Rossman (1984, Herpetologica 40: 388–405) analyzed skull morphology and showed substantial divergence between F. a. abacura and F. a. reinwardtii.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

f. abacura reinwardtii Western Mudsnake

Cundall and Rossman (1984, Herpetologica 40: 388–405) analyzed skull morphology and showed substantial divergence between F. a. abacura and F. a. reinwardtii.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

f. erytrogramma Rainbow Snake

f. erytrogramma erytrogramma Common Rainbow Snake

f. erytrogramma seminola Southern Florida Rainbow Snake

f. streckeri Tamaulipan Hook-Nosed Snake

The previous Standard English names of Ficimia and Gyalopion were misleading relative to their geographic ranges. All are distributed in Mexico, but Ficimia had the moniker “Mexican,” whereas Gyalopion had the name “Plateau,” yet is clearly not confined to any plateau. Given that Ficimia has the easternmost distribution, we call it “Eastern” and call Gyalopion “Western.”

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

g. canum Chihuahuan Hook-Nosed Snake

The previous Standard English names of Ficimia and Gyalopion were misleading relative to their geographic ranges. All are distributed in Mexico, but Ficimia had the moniker “Mexican,” whereas Gyalopion had the name “Plateau,” yet is clearly not confined to any plateau. Given that Ficimia has the easternmost distribution, we call it “Eastern” and call Gyalopion “Western.”

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

g. quadrangulare Thornscrub Hook-Nosed Snake

The previous Standard English names of Ficimia and Gyalopion were misleading relative to their geographic ranges. All are distributed in Mexico, but Ficimia had the moniker “Mexican,” whereas Gyalopion had the name “Plateau,” yet is clearly not confined to any plateau. Given that Ficimia has the easternmost distribution, we call it “Eastern” and call Gyalopion “Western.”

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. striatula Rough Earthsnake

McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) found that Virginia is polyphyletic based on a multi-locus nuclear dataset, and resurrected Haldea for V. striatula.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. gloydi Dusty Hog-Nosed Snake

Werler and Dixon (2000, Texas Snakes, University of Texas Press, Austin) regarded H. n. gloydi to be an allopatric, diagnosable taxon restricted to the low plains - eastern forest ecotone of eastern Texas. Smith et al. (2003, J. Kansas Herpetol. 5: 17–20) followed the taxonomy of Walley and Eckerman (1999, Cat Am. Amph. Rept. 698.1) and did not recognize gloydi.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. kennerlyi Mexican Hog-Nosed Snake

Smith et al. (2003, J. Kansas Herpetol. 5: 17–20), based on two scale characters, separated H. n. kennerlyi from H. n. nasicus and elevated the former to species.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. nasicus Plains Hog-Nosed Snake

Because the three subspecies of H. nasicus have been elevated to species, their respective standard English names remain associated with each. Hence, there is no longer a “Western Hog-nosed Snake.”

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. platirhinos Eastern Hog-Nosed Snake

h. simus Southern Hog-Nosed Snake

h. platurus Yellow-Bellied Seasnake

A recent study (Sanders et al., 2013, Mol. Phylogenet. Evol. 66: 575–591) corrected the long-noted non-monophyly of most sea snake genera (including Pelamis) by recognizing a single large genus Hydrophis, including H. platurus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. chlorophaea Desert Nightsnake

Taxonomy of Hypsiglena has received some critical review since Tanner's revision of the genus (1944, Great Basin Nat. 5: 25–92). Dixon (1965, Southwest. Nat. 10: 125–131) and Dixon and Dean (1986, Southwest. Nat. 31: 307–318) studied a morphological contact zone between northern and southern taxa at the Sonora¬–Sinaloa border in Mexico, finding that it comprised a narrow zone of hybridization with some taxa existing in sympatry. Hardy and McDiarmid (1969, Univ. Kansas Pub. Mus. Nat. Hist. 18: 39–252) examined specimens across the range of this presumptive contact and elsewhere in western Mexico and concluded that no morphological characters existed to separate torquata and ochrorhyncha, except maybe nuchal patterns, which they decided (p. 170) was "a case of pattern dimorphism in a single, otherwise uniform, species." Grismer et al. (1994, Bull. So. California Acad. Sci. 93: 45–80) dismissed the recognition of subspecies in Baja California, stating, without evidence, that the subspecies intergrade widely. Mulcahy (2008,Mol. Phylogenet. Evol. 46: 1095–1115) conducted a comprehensive phylogeographic study of Hypsiglena based on an mtDNA analysis of >150 individuals. Mulcahy (2008) recognized six species in what was considered H. torquata, five of which are consistent with previously described lineages (e.g., subspecies), while one represents a unique lineage that remains to be described. Mulcahy (2008) also recommended maintaining the subspecies designations for several of the widespread, polymorphic species, which may represent incipient species. The nominal species H. torquata is now restricted to Mexico, three described forms occur in the USA, and the undescribed form is endemic to the Cochise Filter Barrier area of southeastern Arizona and associated New Mexico.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. chlorophaea chlorophaea Sonoran Nightsnake

h. chlorophaea deserticola Northern Desert Nightsnake

h. chlorophaea loreala Mesa Verde Nightsnake

h. jani Chihuahuan Nightsnake

Taxonomy of Hypsiglena has received some critical review since Tanner's revision of the genus (1944, Great Basin Nat. 5: 25–92). Dixon (1965, Southwest. Nat. 10: 125–131) and Dixon and Dean (1986, Southwest. Nat. 31: 307–318) studied a morphological contact zone between northern and southern taxa at the Sonora¬–Sinaloa border in Mexico, finding that it comprised a narrow zone of hybridization with some taxa existing in sympatry. Hardy and McDiarmid (1969, Univ. Kansas Pub. Mus. Nat. Hist. 18: 39–252) examined specimens across the range of this presumptive contact and elsewhere in western Mexico and concluded that no morphological characters existed to separate torquata and ochrorhyncha, except maybe nuchal patterns, which they decided (p. 170) was "a case of pattern dimorphism in a single, otherwise uniform, species." Grismer et al. (1994, Bull. So. California Acad. Sci. 93: 45–80) dismissed the recognition of subspecies in Baja California, stating, without evidence, that the subspecies intergrade widely. Mulcahy (2008,Mol. Phylogenet. Evol. 46: 1095–1115) conducted a comprehensive phylogeographic study of Hypsiglena based on an mtDNA analysis of >150 individuals. Mulcahy (2008) recognized six species in what was considered H. torquata, five of which are consistent with previously described lineages (e.g., subspecies), while one represents a unique lineage that remains to be described. Mulcahy (2008) also recommended maintaining the subspecies designations for several of the widespread, polymorphic species, which may represent incipient species. The nominal species H. torquata is now restricted to Mexico, three described forms occur in the USA, and the undescribed form is endemic to the Cochise Filter Barrier area of southeastern Arizona and associated New Mexico.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. jani texana Texas Nightsnake

h. ochrorhyncha Coast Nightsnake

Taxonomy of Hypsiglena has received some critical review since Tanner's revision of the genus (1944, Great Basin Nat. 5: 25–92). Dixon (1965, Southwest. Nat. 10: 125–131) and Dixon and Dean (1986, Southwest. Nat. 31: 307–318) studied a morphological contact zone between northern and southern taxa at the Sonora¬–Sinaloa border in Mexico, finding that it comprised a narrow zone of hybridization with some taxa existing in sympatry. Hardy and McDiarmid (1969, Univ. Kansas Pub. Mus. Nat. Hist. 18: 39–252) examined specimens across the range of this presumptive contact and elsewhere in western Mexico and concluded that no morphological characters existed to separate torquata and ochrorhyncha, except maybe nuchal patterns, which they decided (p. 170) was "a case of pattern dimorphism in a single, otherwise uniform, species." Grismer et al. (1994, Bull. So. California Acad. Sci. 93: 45–80) dismissed the recognition of subspecies in Baja California, stating, without evidence, that the subspecies intergrade widely. Mulcahy (2008,Mol. Phylogenet. Evol. 46: 1095–1115) conducted a comprehensive phylogeographic study of Hypsiglena based on an mtDNA analysis of >150 individuals. Mulcahy (2008) recognized six species in what was considered H. torquata, five of which are consistent with previously described lineages (e.g., subspecies), while one represents a unique lineage that remains to be described. Mulcahy (2008) also recommended maintaining the subspecies designations for several of the widespread, polymorphic species, which may represent incipient species. The nominal species H. torquata is now restricted to Mexico, three described forms occur in the USA, and the undescribed form is endemic to the Cochise Filter Barrier area of southeastern Arizona and associated New Mexico.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

h. ochrorhyncha klauberi San Diego Nightsnake

h. ochrorhyncha nuchalata California Nightsnake

i. braminus Brahminy Blindsnake

The Brahminy Blindsnake is likely native to South Asia, has been reported from 13 states, and is established in Alabama, Arizona, California, Florida, Georgia, Hawaii, Louisiana, Massachusetts, Texas, and Virginia.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-01-13

l. alterna Gray-Banded Kingsnake

Garstka (1982, Breviora 466: 1–35) and more recently Bryson et al. (2007, Mol. Phylogenet. Evol. 43: 674–684) reviewed the mexicana species group of Lampropeltis. Based on the more recent molecular work, it appears that the recognition of the traditional species of alterna, mexicana and triangulum may be incorrect. Until more data are available to resolve the taxonomy of these groups, we withhold making any changes. Given the apparent complexity of the situation and the widespread morphological variation of L. alterna, we do not recognize any subspecies, though Hilken and Schlepper (1998, Salamandra 34: 97–124) argued for recognition of L. alterna alterna and L. a. blairi. Recent work by Ruane et al. (2014, Syst. Biol. 63: 231–250) showed that this is an evolutionarily distinct lineage, and clearly represents a separate species. Previous work showing affinity with L. triangulum based on mitochondrial data was misled by an apparent genome capture.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. annulata Mexican Milksnake

This species comprises a primarily Mexican lineage of the former L. triangulum, and is of uncertain occurrence in the United States, possibly along the Rio Grande in southern Texas (Ruane et al. 2014, Syst. Biol. 63: 231–250).

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. californiae California Kingsnake

Previously considered a subspecies of L. getula, Pyron and Burbrink (2009, Mol. Ecol. 18: 2443–3457 and 2009, Zootaxa 2241: 22–32), demonstrated that this is a distinct species.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. calligaster Prairie Kingsnake

McKelvy and Burbrink (2016. Mol. Phylogenet. Evol. 106: 61–72), using multilocus sequence data, found L. rhombomaculata and L. occipitolineata distinct from L. calligaster.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

l. elapsoides Scarlet Kingsnake

Using multiple nuclear and mitochondrial genes, Pyron and Burbrink (2009, Mol. Phylogenet. Evol. 52: 524–529) found that L. elapsoides is distinct from L. triangulum. This was confirmed in larger multilocus study with many individuals sampled (Ruane et al. 2014, Syst. Biol. 63: 231–250).

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. extenuata Short-Tailed Kingsnake

Dowling and Maxson (1990, J. Zool. London 221: 77–85), using immunological distance data, found Stilosoma to fall within Lampropeltis. Keogh (1996, Herpetologica 52: 406–416), however, found Stilosoma to be part of the probable sister group to Lampropeltis. Rodriguez-Robles and de Jesus Escobar (1999, Biol. J. Linn. Soc. 68: 355–385) and Bryson et al. (2007, Mol. Phylogenet. Evol. 43: 674–684) corroborated Dowling and Maxson using mtDNA evidence, and demonstrated that recognition of Stilosoma as a genus renders Lampropeltis paraphyletic. This was confirmed and ameliorated in Pyron and Burbrink (2009, Mol. Phylogenet. Evol. 52: 524–529) and confirmed in Ruane et al. (2014, Syst. Biol. 63: 231–250).

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. floridana Florida Kingsnake

Krysko et al. (2017, J. Hered. doi:10.1093/jhered/esw086) used mtDNA and nDNA and inferred three monophyletic groups that generally corresponded to the geographic regions Florida Peninsula, Atlantic Coast, and Eastern Apalachicola Lowlands which contained previously recognized subspecies L. g. floridana, L. g. getula, and L. g. meansi. The authors considered the members of the clades as morphologically diagnosable and elevated them to species. Krysko et al. (op. cit.) also recognized L. nigrita from Mexico and southeastern Arizona.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

l. gentilis Western Milksnake

This species comprises the formerly recognized subspecies L. t. celaenops, L. t. multistriata, L. t. taylori, L. t. amaura (part), L. t. syspila (part), and L. t. annulata (part) (Ruane et al. 2014, Syst. Biol. 63: 231–250).

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. getula Eastern Kingsnake

Krysko et al. (2017, J. Hered. doi:10.1093/jhered/esw086) used mtDNA and nDNA and inferred three monophyletic groups that generally corresponded to the geographic regions Florida Peninsula, Atlantic Coast, and Eastern Apalachicola Lowlands which contained previously recognized subspecies L. g. floridana, L. g. getula, and L. g. meansi. The authors considered the members of the clades as morphologically diagnosable and elevated them to species. Krysko et al. also recognized L. nigrita from Mexico and southeastern Arizona.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. holbrooki Speckled Kingsnake

Formerly considered a subspecies of L. getula, Pyron and Burbrink (2009, Mol. Ecol. 18: 2443–3457 and 2009, Zootaxa 2241: 22–32), demonstrated that this is a distinct species. However, compared to the range of the former subspecies, this taxon occurs only west of the Mississippi River.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. knoblochi Madrean Mountain Kingsnake

Formerly considered a subspecies of L. pyromelana, Burbrink et al. (2011, Mol. Phylogenet. Evol. 60: 445–454) demonstrated the existence of two species using coalescent species delimitation methods and ecological niche modeling. The complex comprises a northern species on the Colorado Plateau (L. pyromelana) and a southern species (L. knoblochi) found primarily on the Sierra Madre Occidental and associated Madrean Sky Islands.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. meansi Apalachicola Kingsnake

Krysko et al. (2017, J. Hered. doi:10.1093/jhered/esw086) used mtDNA and nDNA and inferred three monophyletic groups that generally corresponded to the geographic regions Florida Peninsula, Atlantic Coast, and Eastern Apalachicola Lowlands which contained previously recognized subspecies L. g. floridana, L. g. getula, and L. g. meansi. The authors considered the members of the clades as morphologically diagnosable and elevated them to species. Krysko et al. also recognized L. nigrita from Mexico and southeastern Arizona.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

l. multifasciata Coast Mountain Kingsnake

See entry under L. zonata. This species comprises the formerly recognized subspecies L. z. multifasciata, and includes populations from the Transverse and Coastal ranges south.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. nigra Eastern Black Kingsnake

Formerly considered a subspecies of L. getula, Pyron and Burbrink (2009, Mol. Ecol. 18: 2443–3457 and 2009, Zootaxa 2241:22–32), demonstrated that this is a distinct species.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. nigrita Western Black Kingsnake

Krysko et al. (2017, J. Hered. doi:10.1093/jhered/esw086) used mtDNA and nDNA and inferred three monophyletic groups that generally corresponded to the geographic regions Florida Peninsula, Atlantic Coast, and Eastern Apalachicola Lowlands which contained previously recognized subspecies L. g. floridana, L. g. getula, and L. g. meansi. The authors considered the members of the clades as morphologically diagnosable and elevated them to species. Krysko et al. also recognized L. nigrita from Mexico and southeastern Arizona.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

l. occipitolineata South Florida Mole Kingsnake

McKelvy and Burbrink (2016. Mol. Phylogenet. Evol. 106: 61–72), using multilocus sequence data, found L. rhombomaculata and L. occipitolineata distinct from L. calligaster.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

l. pyromelana Arizona Mountain Kingsnake

Burbrink et al. (2011, Mol. Phylogenet. Evol. 60: 445–454) demonstrated that this species is distinct from L. knoblochi.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. rhombomaculata Northern Mole Kingsnake

McKelvy and Burbrink (2016. Mol. Phylogenet. Evol. 106: 61–72), using multilocus sequence data, found L. rhombomaculata and L. occipitolineata distinct from L. calligaster.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

l. splendida Desert Kingsnake

Formerly considered a subspecies of L. getula, Pyron and Burbrink (2009, Mol. Ecol. 18: 2443–3457 and 2009, Zootaxa 2241: 22–32), demonstrated that this is a distinct species.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. triangulum Eastern Milksnake

Ruane et al. (2014, Syst. Biol. 63: 231–250) used a multi-locus nuclear dataset to show that L. triangulum was polyphyletic as previously recognized, consisting of at least three unrelated species groups. As currently defined, L. triangulum primarily comprises populations of the former subspecies L. t. triangulum, L. t. syspila (part), and L. t. amaura (part).

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. zonata California Mountain Kingsnake

This species was investigated using a multi-locus nuclear dataset (Myers et al., 2013, Mol. Ecol. 21: 5418–5429), finding multiple species-level taxa. This species comprises the formerly recognized subspecies L. z. zonata, L. z. multicincta, and L. z. multifasciata (part), including populations from the Sierra Nevada north.

Note on genus: The composition of this group was recently investigated by Ruane et al. (2014, Syst. Biol. 63: 231–250) and, the traditionally recognized species within this genus were found to represent a monophyletic group. However, the composition of various species has changed substantially.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. septentrionalis Northern Cat-Eyed Snake

The genus Leptodeira and the L. septentrionalis/annulata complex in particular, were investigated using a mitochondrial dataset with rangewide sampling (Daza et al. 2009, Mol. Phylogenet. Evol. 53: 653–657). Those authors found that the latter two species are polyphyletic, and that complex geographic structure exists which does not correspond with the current taxonomy.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

l. orcutti Rosy Boa

See annotation under Charina. Wood et al. (2008, Mol. Phylogenet. Evol. 46: 484–582), used mtDNA and found three main clades within trivirgata that do not correspond to currently recognized subspecies. They concluded that these clades corresponded to two species, L. trivirgata and L. orcutti.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. trivirgata Three-Lined Boa

See annotation under Charina. Wood et al. (2008, Mol. Phylogenet. Evol. 46: 484–582), used mtDNA and found three main clades within trivirgata that do not correspond to currently recognized subspecies. They concluded that these clades corresponded to two species, L. trivirgata and L. orcutti.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. alleni Striped Swampsnake

Note on genus: Using a multi-locus nuclear dataset, McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) found that Regina rigida and R. alleni formed a separate species group containing Seminatrix (which has been found previously), and resurrected Liodytes for these species.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. pygaea Black Swampsnake

Using a multi-locus nuclear dataset, McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) found that Regina rigida and R. alleni formed a separate species group containing Seminatrix (which has been found previously), and resurrected Liodytes for these species.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. pygaea cyclas Southern Florida Swampsnake

l. pygaea paludis Carolina Swampsnake

l. pygaea pygaea Northern Florida Swampsnake

l. rigida Glossy Swampsnake

Using a multi-locus nuclear dataset, McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) found that Regina rigida and R. alleni formed a separate species group containing Seminatrix (which has been found previously), and resurrected Liodytes for these species.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

l. rigida deltae Delta Swampsnake

l. rigida rigida Eastern Glossy Swampsnake

l. rigida sinicola Gulf Swampsnake

m. euryxanthus Sonoran Coralsnake

Slowinski (1995, J. Herpetol. 29: 325–338) presented morphological and biochemical data supporting separation of the genera Micrurus and Micruroides. Castoe et al. (2007, Zoo. J. Linn. Soc. 151:809–831) found that Micruroides was the sister taxon to the remainder of the sampled New World Micrurus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

m. euryxanthus euryxanthus Arizona Coralsnake

m. fulvius Harlequin Coralsnake

Although Castoe et al. and J. Boundy (2006, Joint Meeting Ichthyologists Herpetologists abstracts) presented molecular and morphological evidence, respectively, that M. fulvius and M. tener are distinct species, these data have not been published. However, M. tener has been diagnosed by Campbell and Lamar (2004, in J. A. Campbell and W. W. Lamar [eds.], Venomous Reptiles of the Western Hemisphere, Comstock, Publ. Assoc., Ithaca, Pp. 195–197). Using over 1,097 microsatellites, Castoe et al. (2012, Molec. Ecol. Resources 12: 1105–1113) demonstrated that M. fulvius (east of the Mississippi River) is distinct (not sharing genes) with M. tener, which cannot be differentiated from Mexican populations of M. bernardi and M. tamaulipensis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

m. tener Texas Coralsnake

Although Castoe et al. and J. Boundy (2006, Joint Meeting Ichthyologists Herpetologists abstracts) presented molecular and morphological evidence, respectively, that M. fulvius and M. tener are distinct species, these data have not been published. However, this species has been diagnosed by Campbell and Lamar (2004, in J. A. Campbell and W. W. Lamar [eds.], Venomous Reptiles of the Western Hemisphere, Comstock, Publ. Assoc., Ithaca, Pp. 195–197). Using over 1,097 microsatellites, Castoe et al. (2012, Molec. Ecol. Resources 12: 1105–1113) demonstrated that M. fulvius (east of the Mississippi River) is distinct (not sharing genes) with M. tener, which cannot be differentiated from Mexican populations of M. bernardi and M. tamaulipensis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

m. tener tener Texas Gulf-Coast Coralsnake

n. clarkii Saltmarsh Snake

Lawson et al. (1991, Copeia 1991: 638–659) presented allozyme data that supported the separation of clarkii and fasciata.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. clarkii clarkii Gulf Saltmarsh Watersnake

n. clarkii compressicauda Mangrove Saltmarsh Watersnake

n. clarkii taeniata Atlantic Saltmarsh Watersnake

Dunson (1979, Florida Scientist 42: 102–112) synonymized N. c. taeniata with N. c. compressicauda, concluding that it was pattern variant of the latter. Lawson et al. (1991, Copeia 1991: 638–659) resurrected N. c. taeniata on the basis of allozyme data, although the genetic distances were minute.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. cyclopion Mississippi Green Watersnake

n. erythrogaster Plain-Bellied Watersnake

Makowsky et al. (2010, Mol. Phylogenet. Evol.55: 985–995) demonstrated using mitochondrial data that this taxon represents a single widespread species with no concordance to any of the described subspecies. As such we do not recognize subspecies.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. fasciata Southern Watersnake

Allozyme data indicate that N. fasciata forms two clades, differentiated on the mid-Florida Panhandle (Lawson et al., 1991, Copeia 1991: 638–659). Also see note under N. sipedon.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. fasciata confluens Broad-Banded Watersnake

n. fasciata fasciata Banded Watersnake

n. fasciata pictiventris Florida Watersnake

n. floridana Florida Green Watersnake

Elevation of N. floridana from a race of N. cyclopion is supported by data from Pearson (1966, Bull. Serol. Mus. 36: 8), Lawson (1987, J. Herpetol. 21: 140–157), and Sanderson (1993, Brimleyana 19: 83–94). The disjunct populations of floridana were examined by Thompson and Crother (1998, Copeia 1998: 715–719) with allozyme data that revealed no evidence of differentiation.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. harteri Brazos River Watersnake

n. paucimaculata Concho Watersnake

Suggested to be separated from harteri by Rose and Selcer (1989, J. Herpetol. 23: 261–266) and supported by molecular data in Densmore et al. (1992, Herpetologica 48: 60–68).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. rhombifer Diamond-Backed Watersnake

Brandley et al. (2010, Mol. Phylogenet. Evol. 57: 552–560) found evidence for multiple lineages of N. rhombifer. Two lineages were found roughly east and west of the Mississippi River, with a third in Mexico, corresponding to N. r. werleri. However, Brandley et al. sampled only one specimen of the two Mexican subspecies, and it revealed as sister taxon to the U.S. specimens. It cannot be concluded that the Mexican forms are not distinct evolutionary units, and the authors do not conclude otherwise.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

n. rhombifer rhombifer Northern Diamond-Backed Watersnake

Brandley et al. (2010, Mol. Phylogenet. Evol. 57: 552–560) found evidence for multiple lineages of N. rhombifer. Two lineages were found roughly east and west of the Mississippi River, with a third in Mexico, corresponding to N. r. werleri. However, Brandley et al. sampled only one specimen of the two Mexican subspecies, and it revealed as sister taxon to the U.S. specimens. It cannot be concluded that the Mexican forms are not distinct evolutionary units, and the authors do not conclude otherwise.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

n. sipedon Common Watersnake

Numerous examples exist of hybridization between sipedon and fasciata (Conant, 1963, Am. Mus. Novit. 2122: 1–38; Blaney and Blaney, 1979, Herpetologica 35: 350–359; Schwaner et al., 1980, Isozyme Bull. 12: 102; Schwaner and Mount, 1976, Occas. Pap. Mus. Nat. Hist. Univ. Kansas 45: 1–44), and sipedon and fasciata are apparently sister taxa (Pyron et al. 2013, BMC Evol. Biol. 13: 93, doi:10.1186/1471-2148-13-93).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

n. sipedon insularum Lake Erie Watersnake

n. sipedon pleuralis Midland Watersnake

n. sipedon sipedon Northern Watersnake

n. sipedon williamengelsi Carolina Watersnake

n. taxispilota Brown Watersnake

o. aestivus Rough Greensnake

Recognition of the Florida peninsular form described by Grobman (1984, Bull. Florida St. Mus. Biol. Sci. 29: 153–170) is supported by Plummer (1987, Copeia 1987: 483–485). Reviewed by Walley and Plummer (2000, Cat. Am. Amph. Rept. 718).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

o. aestivus aestivus Northern Rough Greensnake

o. aestivus carinatus Florida Rough Greensnake

o. vernalis Smooth Greensnake

Given that Liochlorophis (Oldham and Smith, 1991, Bull. Maryland Herpetol. Soc. 27: 201–215) is the monotypic sister genus to the monotypic genus Opheodrys, recognition of the former taxon is unnecessary, and reduces the amount of information conveyed by the names. As such, we retain vernalis in Opheodrys. The several subspecies described by Grobman (1941, Misc. Pub. Mus. Zool. Univ. Michigan 50: 1–38; 1992, J. Herpetol. 26: 176–186) are based on character clines and not widely recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

o. aeneus Brown Vinesnake

p. alleghaniensis Eastern Ratsnake

Based on the congruence of morphological (Burbrink, 2001, Herpetol. Monogr. 15: 1–53) and mitochondrial data (Burbrink et al., 2000, Evolution 54: 2107–2118), Burbrink divided P. obsoletus into three species (P. alleghaniensis, P. obsoletus and P. spiloides) with no subspecies.

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. bairdi Baird's Ratsnake

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. emoryi Great Plains Ratsnake

Using mitochondrial data, Burbrink (2002, Mol. Phylogenet. Evol. 25: 465–476) found P. guttatus to comprise three distinct lineages, which were elevated to species level. The name P. guttatus was restricted to populations east of the Mississippi River. The populations in western Louisiana and eastern Texas were named P. slowinskii. The subspecies P. g. meahllmorum was not found to be a distinct lineage, and was synonymized with P. emoryi.

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. guttatus Red Cornsnake

Using mitochondrial data, Burbrink (2002, Mol. Phylogenet. Evol. 25: 465–476) found P. guttatus to comprise three distinct lineages, which were elevated to species level. The name P. guttatus was restricted to populations east of the Mississippi River. The populations in western Louisiana and eastern Texas were named P. slowinskii. The subspecies P. g. meahllmorum was not found to be a distinct lineage, and was synonymized with P. emoryi.

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. obsoletus Western Ratsnake

Based on the congruence of morphological (Burbrink, 2001, Herpetol. Monogr. 15: 1–53) and mitochondrial data (Burbrink et al., 2000, Evolution 54: 2107–2118), Burbrink divided P. obsoletus into three species (P. alleghaniensis, P. obsoletus and P. spiloides) with no subspecies.

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. ramspotti Western Foxsnake

Conant (1940, Herpetologica 2: 2) recognized two forms of foxsnakes, one on each side of a geographic disjunction (basically all of Michigan and parts of Indiana and Ohio) with the western form as Pantherophis vulpinus vulpinus and the eastern form as P. v. gloydi. Collins (1991, Herpetol. Rev. 22: 42–43) elevated gloydi to specific status because of its geographic disjunction from vulpinus and the characters noted by Conant (1940, Herpetologica 2: 2). Crother et al. (2011, ISRN Zoology, doi:10.5402/2011/436049) supported the concept of two species, but discovered that the species boundary was the Mississippi River and not the disjunction. The type locality of P. vulpinus is east of the Mississippi River and thus the appropriate available name for the eastern form, leaving the western form unnamed. An interesting side note is that faster evolving microsatellite data reveal a population level separation associated with the geographic hiatus (Row et al., 2011, J Evol. Biol. 24: 2364–2377).

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-10

p. slowinskii Slowinski's Cornsnake

Using mitochondrial data, Burbrink (2002, Mol. Phylogenet. Evol. 25: 465–476) found P. guttatus to comprise three distinct lineages, which were elevated to species level. The name P. guttatus was restricted to populations east of the Mississippi River. The populations in western Louisiana and eastern Texas were named P. slowinskii.

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. spiloides Gray Ratsnake

Based on the congruence of morphological (Burbrink, 2001, Herpetol. Monogr. 15: 1–53) and mitochondrial data (Burbrink et al., 2000, Evolution 54: 2107–2118), Burbrink divided P. obsoletus into three species (P. alleghaniensis, P. obsoletus and P. spiloides) with no subspecies.

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. vulpinus Eastern Foxsnake

Conant (1940, Herpetologica 2: 2) recognized two forms of foxsnakes, one on each side of a geographic disjunction (basically all of Michigan and parts of Indiana and Ohio) with the western form as Pantherophis vulpinus vulpinus and the eastern form as P. v. gloydi. Collins (1991, Herpetol. Rev. 22: 42–43) elevated gloydi to specific status because of its geographic disjunction from vulpinus and the characters noted by Conant (1940, Herpetologica 2: 2). Crother et al. (2011, ISRN Zoology, doi:10.5402/2011/436049) supported the concept of two species, but discovered that the species boundary was the Mississippi River and not the disjunction. The type locality of P. vulpinus is east of the Mississippi River and thus the appropriate available name for the eastern form, leaving the western form unnamed. An interesting side note is that faster evolving microsatellite data reveal a population level separation associated with the geographic hiatus (Row et al., 2011, J Evol. Biol. 24: 2364–2377).

Notes on the genus: Utiger et al. (2002, Russian J. Herpetol. 9: 105–124), using molecular data, divided Elaphe into eight genera. New World Elaphe are part of a clade distinct from Old World species, for which Pantherophis Fitzinger, 1843, was resurrected as the oldest available name. While further splitting of Pantherophis has been proposed (Collins and Taggart, 2008; J. Kansas Herp. 26: 16–18), the use of Pantherophis has helped stabilize the classification of New World ratsnakes for nearly a decade. Thus, we refrain from further division of the genus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

p. browni Saddled Leaf-Nosed Snake

p. decurtatus Spotted Leaf-Nosed Snake

McDiarmid and McCleary (1993, Cat. Am. Amph. Rept.: 579.1–5), argued that the four subspecies of P. browni and five subspecies of P. decurtatus not be recognized. Gardner and Mendelson (2004, J. Herpetol. 38: 187–196), based on morphological data, also concluded that subspecies of P. decurtatus should not be recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. catenifer Gophersnake

Rodriguez-Robles et al. (2000, Mol. Phylogenet. Evol. 14: 35–50) discovered significant internal structuring among P. catenifer populations using mitochondrial data, which may signify the existence of additional species, though they did not attempt reclassification. Pending further study, we retain the present subspecific designations for the group.

Notes on the genus: Using mitochondrial data, Rodríguez-Robles and Jesus-Escobar (2000, Mol. Phylogenet. Evol. 14: 35–50) corroborated the current classification of United States Pituophis into three species: melanoleucus, catenifer, and ruthveni. However, the recognition of ruthveni rendered catenifer paraphyletic, and P. catenifer and P. melanoleucus have geographic structure that does not correspond with currently recognized subspecies. Thus, given further study of this group, some species of Pituophis may undergo taxonomic revision in the near future.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. catenifer affinis Sonoran Gophersnake

p. catenifer annectens San Diego Gophersnake

p. catenifer catenifer Pacific Gophersnake

p. catenifer deserticola Great Basin Gophersnake

p. catenifer pumilus Santa Cruz Island Gophersnake

p. catenifer sayi Bullsnake

p. melanoleucus Eastern Pinesnake

Notes on the genus: Using mitochondrial data, Rodríguez-Robles and Jesus-Escobar (2000, Mol. Phylogenet. Evol. 14: 35–50) corroborated the current classification of United States Pituophis into three species: melanoleucus, catenifer, and ruthveni. However, the recognition of ruthveni rendered catenifer paraphyletic, and P. catenifer and P. melanoleucus have geographic structure that does not correspond with currently recognized subspecies. Thus, given further study of this group, some species of Pituophis may undergo taxonomic revision in the near future.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

p. melanoleucus lodingi Black Pinesnake

p. melanoleucus melanoleucus Northern Pinesnake

p. melanoleucus mugitus Florida Pinesnake

p. ruthveni Louisiana Pinesnake

Reichling (1995, J. Herpetol. 29: 186–198) concluded that ruthveni is a distinct species. Rodriguez-Robles and de Jesús-Escobar (2000, Mol. Phylogenet. Evol. 14: 35–50) argued for the recognition of P. ruthveni, despite lack of significant or independent differentiation from some populations of P. c. sayi using mitochondrial data.

Notes on the genus: Using mitochondrial data, Rodríguez-Robles and Jesus-Escobar (2000, Mol. Phylogenet. Evol. 14: 35–50) corroborated the current classification of United States Pituophis into three species: melanoleucus, catenifer, and ruthveni. However, the recognition of ruthveni rendered catenifer paraphyletic, and P. catenifer and P. melanoleucus have geographic structure that does not correspond with currently recognized subspecies. Thus, given further study of this group, some species of Pituophis may undergo taxonomic revision in the near future.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-11

p. molurus Indian Python

p. molurus bivittatus Burmese Python

Alien Species:

The Burmese Python is native to South and Southeast Asia, has been reported from seven states, and is established in Florida.

Fred Kraus, 2015-01-19

p. sebae Northern African Rock Python

Alien Species:

The Northern African Rock Python is native to sub-Saharan Africa, has been reported from two states, and is established in Florida.

Fred Kraus, 2015-01-19

r. grahamii Graham's Crawfish Snake

Using a multi-locus nuclear dataset, McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) corroborated Alfaro and Arnold (2001, Mol. Phylogenet. Evol. 21: 408–423) and Lawson (1985, Ph.D. dissertation, Louisiana State University) in finding that Regina is polyphyletic, removing R. rigida and R. alleni to Liodytes. Furthermore, R. grahamii and R. septemvittata do not form a strongly supported monophyletic group. Pyron et al. (2013, BMC Evol. Biol. 2013, 13:93,doi:10.1186/1471-2148-13-93) found R. grahamii and R. septemvittata related to Tropidoclonion. Figueroa et al. (2016, PLoS ONE 11(9): e0161070. doi:10.1371/journal.pone.0161070) found grahamii as sister to Tropidoclonion and septemvittata nested within Nerodia. We await further study before making any additional changes.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

r. septemvittata Queensnake

Using a multi-locus nuclear dataset, McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) corroborated Alfaro and Arnold (2001, Mol. Phylogenet. Evol. 21: 408–423) and Lawson (1985, Ph.D. dissertation, Louisiana State University) in finding that Regina is polyphyletic, removing R. rigida and R. alleni to Liodytes. Furthermore, R. grahamii and R. septemvittata do not form a strongly supported monophyletic group. Pyron et al. (2013, BMC Evol. Biol. 2013, 13:93,doi:10.1186/1471-2148-13-93) found R. grahamii and R. septemvittata related to Tropidoclonion. Figueroa et al. (2016, PLoS ONE 11(9): e0161070. doi:10.1371/journal.pone.0161070) found grahamii as sister to Tropidoclonion and septemvittata nested within Nerodia. We await further study before making any additional changes.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

r. dissecta New Mexico Threadsnake

Dixon and Vaughan (2003, Texas J. Sci. 55: 3–24), using morphological data, elevated R. dulcis dissectus to species status, and diagnosed three subspecies within the nominate race, one of which remains unnamed.

Note on genus: Adalsteinsson et al. (2009, Zootaxa 2224: 1–50) demonstrated that the former genus Leptotyphlops was composed of two large clades each composed Old World or New World taxa. The type for the genus Leptotyphlops is associated with Old World taxa, leaving the clade of North and Central American threadsnakes unnamed. The genus Rena has been restored to this group.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

r. dulcis Texas Threadsnake

Dixon and Vaughan (2003, Texas J. Sci. 55: 3–24), using morphological data, elevated R. d. dissectus to species status, and diagnosed three subspecies within the nominate race, one of which remains unnamed.

Note on genus: Adalsteinsson et al. (2009, Zootaxa 2224: 1–50) demonstrated that the former genus Leptotyphlops was composed of two large clades each composed Old World or New World taxa. The type for the genus Leptotyphlops is associated with Old World taxa, leaving the clade of North and Central American threadsnakes unnamed. The genus Rena has been restored to this group.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

r. dulcis dulcis Plains Threadsnake

r. dulcis rubella South Texas Threadsnake

r. humilis Western Threadsnake

Adalsteinsson et al. (2009, Zootaxa 2224: 1–50) demonstrated that the former genus Leptotyphlops was composed of two large clades each composed Old World or New World taxa. The type for the genus Leptotyphlops is associated with Old World taxa, leaving the clade of North and Central American threadsnakes unnamed. The genus Rena has been restored to this group.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

r. humilis cahuilae Desert Threadsnake

r. humilis humilis Southwestern Threadsnake

r. humilis segrega Trans-Pecos Threadsnake

Wallach et al. (2014. Snakes of the World-A Catalogue of Living and Extinct Species, CRC Press) followed Pinto (2010: unpublished PhD Diss, Universidade Federal do Rio de Janeiro.) in recognizing R. segrega as a species. We await publication of the data before we follow Pinto.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

r. humilis utahensis Utah Threadsnake

r. flavilata Pine Woods Littersnake

r. lecontei Long-Nosed Snake

Manier (2004, Biol. J. Linn. Soc. 83: 65–85), in a detailed morphological analysis, concluded that no subspecies should be recognized.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. grahamiae Eastern Patch-Nosed Snake

s. grahamiae grahamiae Mountain Patch-Nosed Snake

s. grahamiae lineata Texas Patch-Nosed Snake

s. hexalepis Western Patch-Nosed Snake

Recognition of the species S. deserticola was made without justification by Bogert and Degenhardt (1961, Am. Mus. Novit. 2064: 13). Bogert (1985, Snake Syst. Newsl. Nov. no. 3) explained that the usage was based on characters discovered previously (Bogert, 1945, Am. Mus. Novit. 1285: 1–14) and on the absence of any intergrades. Although Bogert may be correct, we await a study to demonstrate it and retain S. h. deserticola as a subspecies of S. hexalepis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

s. hexalepis deserticola Big Bend Patch-Nosed Snake

Recognition of the species S. deserticola was made without justification by Bogert and Degenhardt (1961, Am. Mus. Novit. 2064: 13). Bogert (1985, Snake Syst. Newsl. Nov. no. 3) explained that the usage was based on characters discovered previously (Bogert, 1945, Am. Mus. Novit. 1285: 1–14) and on the absence of any intergrades. Although Bogert may be correct, we await a study to demonstrate it and retain S. h. deserticola as a subspecies of S. hexalepis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. hexalepis hexalepis Desert Patch-Nosed Snake

s. hexalepis mojavensis Mohave Patch-Nosed Snake

The spelling of the standard English name has been changed from “Mojave” to “Mohave” for consistency with other names in the list (see note for Crotalus scutulatus).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. hexalepis virgultea Coast Patch-Nosed Snake

s. triaspis Green Ratsnake

Senticolis is more closely related to the New World tribe Lampropeltini than it is to the Old World genus Elaphe (Keogh 1996, Herpetologica 52: 406–416; Utiger et al., 2002, Russian J. Herpetol. 9: 105–124; Burbrink and Lawson, 2007, Mol. Phylogenet. Evol. 43: 173–189 and Pyron and Burbrink, 2009, Mol. Phylogenet. Evol. 52: 524–529).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. triaspis intermedia Northern Green Ratsnake

s. catenatus Eastern Massasauga

Kubatko et al. (2011, Syst. Biol. 60: 393–409) used a multigene data set to infer two clades among the three previously recognized subspecies. One clade contained the eastern subspecies (S. c. catenatus) and the other clade contained the two western subspecies (S. c. tergeminus and S. c. edwardsii). Kubatko et al. (op. cit.) recommended elevating S. c. catenatus. However, if the recommendation was followed at that time, it would also require elevating S. c. tergeminus and the formation of three new combinations. In addition, Holycross et al. (2008, Copeia, 2008: 421–424) discovered that S. c. tergeminus is actually subsumed by S. c. catenatus because the type locality of catenatus is within the range of tergeminus, and that the name Crotalus massassaugusz Kirtland, 1838 would be the available and valid name for the eastern subspecies. As such, tergeminus was not currently a valid name and if the Kubatko et al. recommendation was followed, the specific epithet for the eastern form would be massassaugus. Crother et al. (2011 Bull. Zool. Nomencl. 68: 271–274) submitted a petition to the ICZN for conservation of the names catenatus and tergeminus. The subsequent opinion by the ICZN (2013 Bull. Zool. Nomencl. 70: 282–283) retained the names S. catenatus and S. tergeminus by designation of neotypes for both species. We follow the recommendation of Kubatko et al. (2011, op. cit.) and elevate tergeminus, leaving no recognized subspecies of catenatus. Also see notes under Crotalus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. miliarius Pygmy Rattlesnake

See notes under Crotalus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

s. miliarius barbouri Dusky Pygmy Rattlesnake

Gloyd (1935, Occ. Papers Mus. Zool. Univ. Michigan 322: 1–7) found S. m. barbouri distinct from the other two races by having the lateral spots in 3 series vs. 1–2 series for the other two.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. miliarius miliarius Carolina Pygmy Rattlesnake

s. miliarius streckeri Western Pygmy Rattlesnake

s. tergeminus Western Massasauga

Kubatko et al. (2011, Syst. Biol. 60: 393–409) found mixed signals and limited support for the separation of the subspecies. Also see notes under Crotalus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

s. tergeminus edwardsii Desert Massasauga

s. tergeminus tergeminus Prairie Massasauga

s. semiannulata Western Groundsnake

s. semiannulata semiannulata Variable Groundsnake

s. semiannulata taylori Southern Texas Groundsnake

s. dekayi Dekay's Brownsnake

Pyron et al. (2016, Zool. J. Linn. Soc. 177: 937–949) used high throughput molecular data to detect eight species-level clades within Storeria, but elected to recognize only four clades (three in the United States) that were corroborated by morphology. They concluded against the recognition of subspecies.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

s. occipitomaculata Red-Bellied Snake

No evidence of separate lineages has been found between the sympatric brown and grey color morphs (Grudzien and Owens, 1991, J. Herpetol. 25: 90–92). Pyron et al. (2016, Zool. J. Linn. Soc. 177: 937–949) used high throughput molecular data to detect eight species-level clades within Storeria, but elected to recognize only four clades (three in the United States) that were corroborated by morphology. They concluded against the recognition of subspecies.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

s. victa Florida Brownsnake

Christman (1980, Bull. Florida St. Mus. 25: 157–256) presented evidence, allopatry with no morphological convergence in proximal populations, to suggest species status for victa. This is supported by genomic sequence data (Pyron et al., 2016, Zool. J. Linn. Soc. 177: 937–949).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

t. atriceps Mexican Black-Headed Snake

t. coronata Southeastern Crowned Snake

t. cucullata Trans-pecos Black-headed Snake

The taxonomic status of T. cucullata and T.diabola has been problematic. They have been alternately synonymized (Degenhardt et al., 1976, Texas J. Sci. 17: 225–234; Hillis and Campbell, 1982, Southwest. Nat. 27: 220–221; Irwin and Collins, 1995, Herpetol. Rev. 26: 47) or elevated to species (Collins, 1991, Herpetol. Rev. 22: 42–43). We follow the most recent proposals from Wilson (1999, Smithsonian Inform. Serv. 122: 1–34) and Dixon et al. (2000, Southwest Nat. 45) who both recognized T. cucullata as a species distinct from T. rubra (extralimital) and synonymized T. diabola with the former.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. gracilis Flat-Headed Snake

t. hobartsmithi Smith's Black-headed Snake

t. nigriceps Plains Black-Headed Snake

t. oolitica Rim Rock Crowned Snake

t. planiceps Western Black-Headed Snake

Cole and Hardy (1981, Bull. Am. Mus. Nat. Hist. 17: 201–284) noted local geographic variation but did not recognize any available subspecies of the many disjunct populations.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. relicta Florida Crowned Snake

t. relicta neilli Central Florida Crowned Snake

t. relicta pamlica Coastal Dunes Crowned Snake

t. relicta relicta Peninsula Crowned Snake

t. wilcoxi Chihuahuan Black-Headed Snake

t. yaquia Yaqui Black-Headed Snake

t. atratus Aquatic Gartersnake

Rossman and Stewart (1987, Occ. Pap. Mus. Zool. Louisiana St. Univ. 63: 1–25) recognized atratus as distinct from T. couchii and recommended against recognizing T. a. aquaticus.

Note on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. atratus atratus Santa Cruz Gartersnake

t. atratus hydrophilus Oregon Gartersnake

t. atratus zaxanthus Diablo Range Gartersnake

t. brachystoma Short-Headed Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. butleri Butler's Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press)

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. couchii Sierra Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press)

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. cyrtopsis Black-Necked Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press)

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. cyrtopsis cyrtopsis Western Black-Necked Gartersnake

t. cyrtopsis ocellatus Eastern Black-Necked Gartersnake

t. elegans Terrestrial Gartersnake

Using mitochondrial data, Bronikowski and Arnold (2001, Copeia 2001: 508–513) identified several clades within T. elegans that did not, in some cases, follow phenotypic subspecies boundaries. Hammerson (1999, Amphibians and Reptiles of Colorado. 2nd ed. University of Colorado Press, Boulder) found phenotypes assignable to T. e. arizonae and T. e. vascotanneri outside of their purported distributions within Colorado, and recommended that the two names be synonymized with T. e. vagrans. Hammerson's data supported similar action for Arizona and New Mexico populations as well (J. Boundy, pers. obs.). Thus, we tentatively retain three subspecies.

Note on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. elegans elegans Mountain Gartersnake

t. elegans terrestris Coast Gartersnake

t. elegans vagrans Wandering Gartersnake

t. eques Mexican Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press)

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. eques megalops Brown Gartersnake

t. gigas Giant Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press)

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. hammondii Two-Striped Gartersnake

The extralimital T. digueti was synonymized with T. hammondi by McGuire and Grismer (1993, Herpetologica 49: 354–365).

Note on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. marcianus Checkered Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. marcianus marcianus Marcy's Checkered Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. ordinoides Northwestern Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. proximus Western Ribbonsnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. proximus diabolicus Arid Land Ribbonsnake

t. proximus orarius Gulf Coast Ribbonsnake

t. proximus proximus Orange-Striped Ribbonsnake

t. proximus rubrilineatus Red-Striped Ribbonsnake

t. radix Plains Gartersnake

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Jonathan A. Campbell, R. Alexander Pyron, 2014-12-12

t. rufipunctatus Narrow-Headed Gartersnake

Based on scale microstructure, Chiasson and Lowe (1989, J. Herpetol. 23: 109–118) suggested this taxon be moved from Thamnophis to Nerodia. De Queiroz and Lawson (1994, Biol. J. Linn. Soc. 53: 209–229) rejected the suggested reallocation, based on their finding that rufipunctatus is nested within Thamnophis.

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. saurita Eastern Ribbonsnake

Kraus and Cameron (2016, Herpetol. Rev. 47: 74–75) corrected the spelling to saurita.

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. saurita nitae Blue-Striped Ribbonsnake

Kraus and Cameron (2016, Herpetol. Rev. 47: 74–75) corrected the spelling to saurita.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

t. saurita sackenii Peninsula Ribbonsnake

Kraus and Cameron (2016, Herpetol. Rev. 47: 74–75) corrected the spelling to saurita.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

t. saurita saurita Common Ribbonsnake

Kraus and Cameron (2016, Herpetol. Rev. 47: 74–75) corrected the spelling to saurita.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

t. saurita septentrionalis Northern Ribbonsnake

Kraus and Cameron (2016, Herpetol. Rev. 47: 74–75) corrected the spelling to saurita.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-01-13

t. sirtalis Common Gartersnake

Analyses of mitochondrial and nuclear data suggest that this species may be composed of multiple independently evolving lineages often not concordant with the subspecific taxonomy (F. Burbrink, pers. comm.).

Notes on the genus: The specific and infraspecific status of Thamnophis is based on Rossman et al. (1996, The Garter Snakes: Evolution and Ecology, Univ. Oklahoma Press).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. sirtalis annectens Texas Gartersnake

t. sirtalis concinnus Red-Spotted Gartersnake

t. sirtalis dorsalis New Mexico Gartersnake

t. sirtalis fitchi Valley Gartersnake

t. sirtalis infernalis California Red-Sided Gartersnake

The International Commission on Zoological Nomenclature (2000, Bull. Zool. Nomencl. 57: 191–192, Opinion 1961) has ruled that the name Coluber infernalis be re-associated with Pacific Coast populations referred to as T. s. concinnus by Crother et al. (2000, Herpetol. Circular 29: 73), as suggested by Boundy and Rossman (1995, Copeia 1995: 236–240).

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. sirtalis pallidulus Maritime Gartersnake

t. sirtalis parietalis Red-Sided Gartersnake

t. sirtalis pickeringii Puget Sound Gartersnake

t. sirtalis semifasciatus Chicago Gartersnake

Benton (1980, Zool. J. Linnaean Soc. 68: 307–323) synonymized T. s. semifasciatus with the nominate race, but Rossman et al. (1996, The Gartersnakes. Evolution and Ecology, Univ. Oklahoma Press) resurrected T. s. semifasciatus.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. sirtalis similis Blue-Striped Gartersnake

t. sirtalis sirtalis Eastern Gartersnake

t. sirtalis tetrataenia San Francisco Gartersnake

Action by the International Commission on Zoological Nomenclature (2000, Bull. Zool. Nomencl. 57: 191–192. Opinion 1961) has retained the name Eutaenia s. tetrataenia for San Francisco Peninsula populations of T. sirtalis.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. lambda Sonoran Lyresnake

Devitt et al. (2008, Copeia 2008: 370–387) recognized six species (three extralimital), including T. lambda and T. lyrophanes based on morphological and mitochondrial data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. lyrophanes California Lyresnake

Devitt et al. (2008, Copeia 2008: 370–387) recognized six species (three extralimital), including T. lambda and T. lyrophanes based on morphological and mitochondrial data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. vilkinsonii Texas Lyresnake

LaDuc and Johnson (2003, Herpetologica 59: 364–374) re–elevated T. vilkinsonii to species status.

Note on genus: Devitt et al. (2008, Copeia 2008: 370–387) recognized six species (three extralimital), including T. lambda and T. lyrophanes based on morphological and mitochondrial data.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

t. lineatum Lined Snake

See comments under Regina.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

v. valeriae Smooth Earthsnake

McVay and Carstens (2013, Mol. Phylogenet. Evol. 68: 425–431) found that Virginia is polyphyletic based on a multi-locus nuclear dataset, and resurrected Haldea for V. striatula.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

v. valeriae elegans Western Smooth Snake

v. valeriae pulchra Mountain Earthsnake

Collins (1991, Herpetol. Rev. 22: 42–43) elevated pulchra to specific status. Because no supporting data, aside from allopatric distribution, were published in his list, we retain V. valeriae pulchra.

Brian I. Crother (Chair), Jeff Boundy, Frank T. Burbrink, Sara Ruane, 2018-02-12

v. valeriae valeriae Eastern Smooth Earthsnake

squamata — lizard

a. picticauda Peters's Rock Agama

Peters’s Rock Agama is native to western Africa and is established in Florida. Earlier confusion about the taxonomy of these lizards (Enge et al. 2004. Florida Scientist 67: 303–310) has been resolved by Nuñez et al. (2016, Bull. Florida Mus. Nat. Hist. 9: 138–146).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-01-13

a. ameiva Giant Ameiva

Alien Species:

The Giant Ameiva is native to Amazonia and is established in Florida. Earlier confusion about the taxonomy of these lizards (Wilson and Porras, 1983, The Ecological Impact of Man on the South Florida Herpetofauna. Univ. Kansas Mus. Nat. Hist. Spec. Publ. 9: 1–89) has been resolved by Ugueto and Harvey (2011, Herpetol. Monogr. 25: 113–170).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. praesignis Dusky Giant Ameiva

Alien Species:

The Dusky Giant Ameiva is native to lower Central America and northwestern South America; it is established in Florida. Earlier confusion about the taxonomy of these lizards (Wilson and Porras, 1983, The Ecological Impact of Man on the South Florida Herpetofauna. Univ. Kansas Mus. Nat. Hist. Spec. Publ. 9: 1–89) has been resolved by Ugueto and Harvey (2011, Herpetol. Monogr. 25: 113–170).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. alexanderae Temblor Legless Lizard

Taxonomy for Anniella follows Papenfuss and Parham (2013, Breviora 536: 1–17), who recognized five species for specimens previously referred to A. pulchra based on molecular and morphological evidence. Some of the common names proposed by Papenfuss and Parham (op. cit.) have been changed in the interest of brevity and descriptive accuracy.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. campi Big Spring Legless Lizard

Taxonomy for Anniella follows Papenfuss and Parham (2013, Breviora 536: 1–17), who recognized five species for specimens previously referred to A. pulchra based on molecular and morphological evidence. Some of the common names proposed by Papenfuss and Parham (op. cit.) have been changed in the interest of brevity and descriptive accuracy.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. grinnelli Bakersfield Legless Lizard

Taxonomy for Anniella follows Papenfuss and Parham (2013, Breviora 536: 1–17), who recognized five species for specimens previously referred to A. pulchra based on molecular and morphological evidence. Some of the common names proposed by Papenfuss and Parham (op. cit.) have been changed in the interest of brevity and descriptive accuracy.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. pulchra Northern Legless Lizard

Taxonomy for Anniella follows Papenfuss and Parham (2013, Breviora 536: 1–17), who recognized five species for specimens previously referred to A. pulchra based on molecular and morphological evidence. Some of the common names proposed by Papenfuss and Parham (op. cit.) have been changed in the interest of brevity and descriptive accuracy.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. stebbinsi San Diegan Legless Lizard

Taxonomy for Anniella follows Papenfuss and Parham (2013, Breviora 536: 1–17), who recognized five species for specimens previously referred to A. pulchra based on molecular and morphological evidence. Some of the common names proposed by Papenfuss and Parham (op. cit.) have been changed in the interest of brevity and descriptive accuracy.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. carolinensis Green Anole

Tollis et al. (2012, PLoS ONE 7(6): e38474) and Campbell-Staton et al. (2012, Ecol. Evol. 2: 2274–2284) provided evidence for the existence of five mutually exclusive mtDNA clades within A. carolinensis. Although nDNA (Tollis et al., op. cit.) corroborated the existence of some of these units, it also suggested the existence of gene flow between others. More extensive geographic sampling by Tollis and Boissinot (2014, Genetica 142: 59–72) revealed that two of the five mtDNA clades are nested but supported five genetic clusters, with admixture. More extensive nDNA sampling by Manthey et al. (2016, Ecol. and Evol. doi: 10.1002/ece3.2547) corroborated those clusters. Because the distributions of the subspecies proposed by Vance (1991, Bull. Maryland Herpetol. Soc. 27: 43–89) do not match those of any of the five genetic clusters, we have not recognized subspecies; however, the existence of fixed differences and an estimated Pliocene divergence between southern Florida and the remaining populations (Manthey et al., 2016, op. cit) suggests that the possibility of more than one species. Species delimitation studies involving contact zones between the phylogeographic groups are needed.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder and Adam D. Leaché, 2018-01-13

a. chlorocyanus Hispaniolan Green Anole

Alien Species: The Hispaniolan Green Anole is native to Hispaniola and is established in Florida

Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press) and modifications by Vance (1991, Bull. Maryland Herpetol. Soc. 27: 43–89; description of A. carolinensis seminolus). Some authors (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116; Nicholson et al., 2012, Zootaxa 3477: 1–108) divide Anolis into several genera and these are included in parentheses (as subclades). Assignments of species covered in this checklist to the genera of Nicholson et al. (2012) is as follows: Anolis (carolinensis, porcatus), Audantia (cybotes), Ctenonotus (cristatellus, distichus, ferreus), Dactyloa (trinitatis), Deiroptyx (equestris), Norops (garmani, sagrei), Xiphosurus (chlorocyanus).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. cristatellus Crested Anole

Alien Species: The Puerto Rican Crested Anole is native to Puerto Rico and the Virgin Islands and is established in Florida. Subspecific identifications have been given for the Dade County specimens by Schwartz and Henderson (1988, Contrib. Biol. Geol. Milwaukee Publ. Mus. 74: 1–264; 1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press).

Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press) and modifications by Vance (1991, Bull. Maryland Herpetol. Soc. 27: 43–89; description of A. carolinensis seminolus). Some authors (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116; Nicholson et al., 2012, Zootaxa 3477: 1–108) divide Anolis into several genera and these are included in parentheses (as subclades). Assignments of species covered in this checklist to the genera of Nicholson et al. (2012) is as follows: Anolis (carolinensis, porcatus), Audantia (cybotes), Ctenonotus (cristatellus, distichus, ferreus), Dactyloa (trinitatis), Deiroptyx (equestris), Norops (garmani, sagrei), Xiphosurus (chlorocyanus).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. cristatellus cristatellus Puerto Rican Crested Anole

Alien Species: The Puerto Rican Crested Anole is native to Puerto Rico and the Virgin Islands and is established in Florida. Subspecific identifications have been given for the Miami-Dade County specimens by Schwartz and Henderson (1988, Contrib. Biol. Geol. Milwaukee Publ. Mus. 74: 1–264; 1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press).

Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press) and modifications by Vance (1991, Bull. Maryland Herpetol. Soc. 27: 43–89; description of A. carolinensis seminolus). Some authors (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116; Nicholson et al., 2012, Zootaxa 3477: 1–108) divide Anolis into several genera and these are included in parentheses (as subclades). Assignments of species covered in this checklist to the genera of Nicholson et al. (2012) is as follows: Anolis (carolinensis, porcatus), Audantia (cybotes), Ctenonotus (cristatellus, distichus, ferreus), Dactyloa (trinitatis), Deiroptyx (equestris), Norops (garmani, sagrei), Xiphosurus (chlorocyanus).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. cybotes Large-Headed Anole

Alien Species: The Large-headed Anole is native to Hispaniola and the Bahamas and is established in Florida.

Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press) and modifications by Vance (1991, Bull. Maryland Herpetol. Soc. 27: 43–89; description of A. carolinensis seminolus). Some authors (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116; Nicholson et al., 2012, Zootaxa 3477: 1–108) divide Anolis into several genera and these are included in parentheses (as subclades). Assignments of species covered in this checklist to the genera of Nicholson et al. (2012) is as follows: Anolis (carolinensis, porcatus), Audantia (cybotes), Ctenonotus (cristatellus, distichus, ferreus), Dactyloa (trinitatis), Deiroptyx (equestris), Norops (garmani, sagrei), Xiphosurus (chlorocyanus).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. cybotes cybotes Common Large-Headed Anole

Alien Species: The Miami-Dade County population has been identified as A. c. cybotes (Schwartz and Henderson, 1988, Contrib. Biol. Geol. Milwaukee Pub. Mus. 74: 1–264). No subspecific identification for the Broward County or Palm Beach County populations has been provided.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. distichus Bark Anole

The potential natural occurrence of A. (Ctenonotus) distichus in Florida is an unresolved issue. Current populations show evidence of hybridization between introduced A. d. dominicensis and another form, but the origin of the other form is currently unknown. Smith and McCauley (1948, Proc. Biol. Soc. Washington 61: 159–166) named it as the subspecies A. d. floridanus based on differences from the Bahamian and Hispaniolan specimens. Schwartz (1968, Bull. Mus. Comp. Zool. 137: 255–310) reviewed morphological variation in A. distichus and confirmed differences between Floridian versus Bahamian and Hispaniolan populations. He considered A. d. floridanus to have colonized Florida recently, either by natural dispersal or human introduction, and that the Bimini chain (A. d. biminiensis) and Andros Island (A. d. distichoides) represented the most likely sources. A detailed study of genetic variation in A. distichus, similar to that done for A. sagrei (Kolbe et al., 2004, Nature 431: 177–181) and including the introduced populations, would help to clarify this issue.

Alien species: The Bark Anole is native to Hispaniola, has been reported from two states, and is established in Florida.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, Fred Kraus, and Kenneth L. Krysko, 2018-02-02

a. distichus dominicensis Green Bark Anole

Alien Species: Anolis distichus dominicensis is established in Miami, Florida (King and Krakauer, 1966, Quart. J. Florida Acad. Sci. 29: 144–154; Wilson and Porras, 1983, Univ. Kansas Mus. Nat. Hist. Spec. Publ. 9: 1–89). Another subspecies, Anolis distichus ignigularis, was introduced to Miami-Dade County, Florida (King and Krakauer, 1966, op.cit. and was listed as occurring there by Schwartz and Henderson (1988, Contrib. Biol. Geol. Milwaukee Pub. Mus. 74: 1–264; 1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press); however, according to Wilson and Porras (1983, Univ. Kansas Mus. Nat. Hist. Spec. Publ. 9: 1–89), this population is no longer extant. Hybridization appears to have occurred between A. d. dominicensis and A. d. floridanus (Miyamoto et al., 1986, Copeia 1986: 76–86; see A. d. floridanus).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. distichus floridanus Florida Bark Anole

The potential natural occurrence of A. (Ctenonotus) distichus in Florida is an unresolved issue. Current populations show evidence of hybridization between introduced A. d. dominicensis and another form, but the origin of the other form is currently unknown. Smith and McCauley (1948, Proc. Biol. Soc. Washington 61: 159–166) named it as the subspecies A. d. floridanus based on differences from the Bahamian and Hispaniolan specimens. Schwartz (1968, Bull. Mus. Comp. Zool. 137: 255–310) reviewed morphological variation in A. distichus and confirmed differences between Floridian versus Bahamian and Hispaniolan populations. He considered A. d. floridanus to have colonized Florida recently, either by natural dispersal or human introduction, and that the Bimini chain (A. d. biminiensis) and Andros Island (A. d. distichoides) represented the most likely sources. A detailed study of genetic variation in A. distichus, similar to that done for A. sagrei (Kolbe et al., 2004, Nature 431: 177–181) and including the introduced populations, would help to clarify this issue.

Alien species: Schwartz (1968, Bull. Mus. Comp. Zool. 137: 255–310) reviewed the evidence and discussed alternative hypotheses concerning the occurrence of Anolis distichus floridanus in Florida and concluded that this taxon was most likely introduced from Andros Island in The Bahamas; nevertheless, Wilson and Porras (1983, Univ. Kansas Mus. Nat. Hist. Spec. Publ. 9: 1–89) considered it a native component of the Florida herpetofauna. Although the specimens of A. d. floridanus examined by Schwartz (1968, Bull. Mus. Comp. Zool. 137: 255–310) are distinguishable from those of A. d. dominicensis, more recent samples of Bark Anoles from Florida form a continuum, suggesting intergradation between the two subspecies (Miyamoto et al., 1986, Copeia 1986: 76–86).

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, Adam D. Leaché, Fred Kraus, and Kenneth L. Krysko, 2018-02-02

a. equestris Knight Anole

Alien Species:

The Knight Anole is native to Cuba and is established in Florida and Hawaii.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. equestris equestris Western Knight Anole

Alien Species:

The subspecific identification for the Florida population was given by Schwartz and Henderson (1988, Contrib. Biol. Geol. Milwaukee Pub. Mus. 74: 1–264; 1991, Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History, University of Florida Press); that for the Hawaiian population was given by Lazell and McKeown (1998, Bull. Chicago Herpetol. Soc. 33: 181).

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. garmani Jamaican Giant Anole

Alien Species: The Jamaican Giant Anole is native to Jamaica and is established in Florida.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. porcatus Cuban Green Anole

Alien Species:

The Cuban Green Anole is native to Cuba, has been reported in Florida, but there is currently no evidence that the species itself is established, although mitochondrial DNA from this species is present in Florida, making it clear that the species had been introduced at one time (Kolby et al., 2007, Conserv. Biol. 21: 1612–1625). The status of this species in Florida, therefore, requires clarification.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. sagrei Brown Anole

Alien Species: The Brown Anole is native to Cuba and the Bahamas, has been reported from 13 states, and is established in Alabama, California, Florida, Georgia, Hawaii, Louisiana, and Texas. Reports from other southern states require confirmation of establishment.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. sagrei sagrei Cuban Brown Anole

Alien Species: According to Conant and Collins (1991, Reptiles and Amphibians of Eastern and Central North America, Houghton Mifflin Co.), two subspecies, A. s. sagrei and A. s. ordinatus were introduced to southern Florida, but they can no longer be distinguished from one another and differ from both original races. Lee (1992, Copeia 1992: 942–954) presented evidence that the Florida populations bear a much stronger phenotypic resemblance to populations from Cuba (A. s. sagrei) than to those from the Bahamas (A. s. ordinatus). Kolbe et al. (2004, Nature 431: 177–181) present evidence for multiple introductions of this species from Cuba to Florida, which suggests that A. s. greyi may also have been involved.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. trinitatis St. Vincent Bush Anole

Alien Species: The St. Vincent Bush Anole is native to St. Vincent, Lesser Antilles, and is established in Florida.

Note on genus: Taxonomy for Anolis follows Williams (1976, Breviora 440: 1–21) with addition of subspecies from Schwartz and Henderson (1991, Amphibians and Reptiles of the West Indies, University of Florida Press) and modi cations described in the notes below. Some authors divide Anolis into five (e.g., Guyer and Savage, 1986, Syst. Zool. 35: 509–531; 1992, Syst. Biol. 41: 89–110; Savage and Guyer, 1989, Amphibia-Reptilia 10: 105–116) or 8 (Nicholson et al., 2012, Zootaxa 3477: 1–108) genera (for criticisms see Williams, 1989, in C. A. Woods [ed.], Biogeography of the West Indies, Sandhill Crane Press :433–477; Cannatella and de Queiroz, 1989, Syst. Zool. 38: 57–69; Jackman et al., 1999, Syst. Biol. 48: 254–285; Poe, 2004, Herpetol. Monogr. 18: 37–89; 2013, Zootaxa 3626: 295–299). Other authors (e.g., Nicholson, 2002, Herpetol. Monogr. 16: 93–120; Brandley and de Queiroz, 2004, Herpetol. Monogr. 18: 90–126; Castañeda and de Queiroz, 2011, Mol. Phylogenet. Evol. 61: 784–800; 2013, Bull. Mus. Comp. Zool. 160: 345–398) use the name Anolis for the larger clade, applying the other names to various of its subclades (sometimes with different circumscriptions than the genera with the same names). We have adopted the second approach and included names of subclades parenthetically, where applicable.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. exsanguis Chihuahuan Spotted Whiptail

Unisexual.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. flagellicauda Gila Spotted Whiptail

Unisexual.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. gularis Common Spotted Whiptail

See comment under A. scarlais.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. gularis gularis Texas Spotted Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. hyperythra Orange-Throated Whiptail

A multivariate analysis of morphological variation in A. hyperythra by Taylor and Walker (2014, Southwest. Nat. 59: 221–227) found evidence of differentiation between populations north and south of the Isthmus of La Paz, which have previously been recognized as the subspecies A. h. beldingi and A. h. hyperythra, respectively (e.g., Wright, 1994, in Brown and Wright [eds.], Herpetology of the North American Deserts, Southwestern Herpetologists Society :255–271). The results of that study did not support the recognition of A. h. schmidti.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. hyperythra beldingi Belding's Orange Throated Whiptail

A multivariate analysis of morphological variation in A. hyperythra by Taylor and Walker (2014, Southwest. Nat. 59: 221–227) found evidence of differentiation between populations north and south of the Isthmus of La Paz, which have previously been recognized as the subspecies A. h. beldingi and A. h. hyperythra, respectively (e.g., Wright, 1994, in Brown and Wright [eds.], Herpetology of the North American Deserts, Southwestern Herpetologists Society :255–271). The results of that study did not support the recognition of A. h. schmidti.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. inornata Little Striped Whiptail

Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157) recognized six subspecies of Aspidoscelis inornata in the United States. Collins (1997, SSAR Herpetol. Circ. 25), treated three of them, arizonae, gypsi, and pai, as separate species (but see note on A. i. gypsi). Walker et al. (1996, J. Herpetol. 30: 271–275) called into question some of the characters used by Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129– 157) to separate Aspidoscelis inornata junipera from A. i. heptagramma. Walker et al. (2012, Herpetol. Conserv. Biol. 7: 265–275) and Sullivan et al. (2013, Copeia 2013:366– 377) provided morphological evidence for the separation of A. pai from A. arizonae; however, Sullivan et al. (op. cit.) found that A. arizonae was not morphologically distinguishable from A. i. llanuras. Sullivan et al. (2014, Copeia 2014: 519–529) found incongruence between mtDNA clades and several of the subspecies of A. inornata. We have followed those authors in no longer recognizing arizonae and pai as species separate from A. inornata and in considering heptagramma, junipera, and llanuras to form a single taxonomic entity. A comprehensive study of gene flow within the Little Striped Whiptail complex, including the Mexican populations, is needed.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. inornata arizonae Arizona Striped Whiptail

Walker et al. (2012, Herpetol. Conserv. Biol. 7: 265–275) and Sullivan et al. (2013, Copeia 2013:366– 377) provided morphological evidence for the separation of A. pai from A. arizonae; however, Sullivan et al. (op. cit.) found that A. arizonae was not morphologically distinguishable from A. i. llanuras. Sullivan et al. (2014, Copeia 2014: 519–529) found incongruence between mtDNA clades and several of the subspecies of A. inornata. We have followed those authors in no longer recognizing arizonae and pai as species separate from A. inornata and in considering heptagramma, junipera, and llanuras to form a single taxonomic entity. A comprehensive study of gene ow within the Little Striped Whiptail complex, including the Mexican populations, is needed.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. inornata gypsi Little White Whiptail

Rosenblum and Harmon (2010, Evolution 65: 946–960), in a study based on nuclear and mitochondrial DNA, coloration, and body size and proportions, concluded that although whiptails from the gypsum sands had diverged more from their dark soil counterparts in terms of body size and shape than sympatric earless and fence lizards (see notes on Holbrookia maculata ruthveni and Sceloporus cowlesi), the genetic data indicate that the whiptails are failing to speciate. This conclusion suggests that it is more appropriate to recognize the taxon not as a species (as proposed by Collins, 1997, SSAR Herpetol. Circ. 25) but as a subspecies of A. inornata.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. inornata heptagramma Northern Striped Whiptail

Walker et al. (2012, Herpetol. Conserv. Biol. 7: 265–275) and Sullivan et al. (2013, Copeia 2013:366– 377) provided morphological evidence for the separation of A. pai from A. arizonae; however, Sullivan et al. (op. cit.) found that A. arizonae was not morphologically distinguishable from A. i. llanuras. Sullivan et al. (2014, Copeia 2014: 519–529) found incongruence between mtDNA clades and several of the subspecies of A. inornata. We have followed those authors in no longer recognizing arizonae and pai as species separate from A. inornata and in considering heptagramma, junipera, and llanuras to form a single taxonomic entity. A comprehensive study of gene ow within the Little Striped Whiptail complex, including the Mexican populations, is needed.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. inornata pai Pai Striped Whiptail

Walker et al. (2012, Herpetol. Conserv. Biol. 7: 265–275) and Sullivan et al. (2013, Copeia 2013:366– 377) provided morphological evidence for the separation of A. pai from A. arizonae; however, Sullivan et al. (op. cit.) found that A. arizonae was not morphologically distinguishable from A. i. llanuras. Sullivan et al. (2014, Copeia 2014: 519–529) found incongruence between mtDNA clades and several of the subspecies of A. inornata. We have followed those authors in no longer recognizing arizonae and pai as species separate from A. inornata and in considering heptagramma, junipera, and llanuras to form a single taxonomic entity. A comprehensive study of gene ow within the Little Striped Whiptail complex, including the Mexican populations, is needed.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. laredoensis Laredo Striped Whiptail

Unisexual. Abuhteba et al. (2001, Copeia 2001: 262–266) interpreted histoincompatibility between the members of two pattern classes within Aspidoscelis laredoensis as evidence for separate hybrid origins of the corresponding clones. The authors noted that two of them are planning to restrict the name A. laredoensis to one of the clones and propose a new species name for the other.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. marmorata Marbled Whiptail

Dessauer and Cole (1991, Copeia 1991: 622–637; see also Dessauer et al., 2000, Bull. Am. Mus. Nat. Hist. 246: 1–148) presented evidence of both differentiation and interbreeding between A. marmorata and A. tigris along a transect near the southern part of the border between Arizona and New Mexico, including a narrow (3 km) hybrid zone in which hybrid indices based on color patterns and allele frequencies changed abruptly in concordant step clines. Although those authors interpreted their data as reflecting incomplete speciation between the two forms (i.e., a single species), the same data can be interpreted alternatively as reflecting largely separate gene pools (i.e., two species). Following the terminology of de Queiroz (1998, in D. J. Howard and S. H. Berlocher [eds.], Endless Forms: Species and Speciation, Oxford University Press, Pp. 57–75), they are here considered incompletely separated species.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. marmorata marmorata Western Marbled Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81). Taylor et al. (2001, Am. Mus. Novit. 3345: 1–65) presented evidence for hybridization between A. tesselata and A. marmorata, but there is no indication that this hybridization has produced a new hybrid species.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. marmorata reticuloriens Eastern Marbled Whiptail

See note on A. tesselata concerning hybridization between that species and A. m. reticuloriens.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. motaguae Giant Whiptail

Alien Species:

The Giant Whiptail is native to Central America and is established in Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

a. neavesi Neaves’ Whiptail

Unisexual. This tetraploid parthenogenetic species of Aspidoscelis was generated in the laboratory by hybridization between A. exsanguis and A. inornata (Lutes et al., 2011, Proc. Natl. Acad. Sci. USA 108: 9910–9915; Cole et al., 2014, Breviora 539: 1–19). It is not known to occur in the wild.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. neomexicana New Mexico Whiptail

Unisexual. Manning et al. (2005, Am. Mus. Novit. 3492: 1–56) presented evidence for hybridization between A. neomexicana and A. sexlineatus viridis, but there is no indication either that this hybridization has produced a new hybrid species or that it is leading to the fusion of the two species.

Note on genus: Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81) with modifications by Camp (1916, Univ. California Pub. Zool. 17: 63–74; proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti), Dessauer and Cole (1991, Copeia 1991: 622–637; recognition of A. marmorata (tigris) reticuloriens), Trauth (1992, Texas J. Sci. 44: 437–443; proposal of A. sexlineata stephensae), Wright and Lowe (1993, J. Arizona-Nevada Acad. Sci. 27: 129–157; proposals of A. inornatus gypsi, A. i. junipera, A. i. llanuras, and A. i. pai), Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae), Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris, and use of the name A. t. punctilinealis for the taxon formerly called A. t. gracilis), Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members), ICZN (1999, Bull. Zool. Nomencl. 56: 162–163) precedence of the name A. neomexicana over A. perplexa; Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61; use of Aspidoscelis for a clade containing all of the whiptail species native to North America), Cole et al. (2014, Breviora 539:1–19; proposal of A. neavesi), and those described in additional notes below. Maslin and Secoy (op. cit.) and Wright (op. cit.) are the sources for information on reproductive mode.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. neotesselata Colorado Checkered Whiptail

Walker et al. (1997, Herpetologica 53: 233–259; restriction of the name A. tesselata to the diploid members of the species formerly referred to by that name and recognition of the species A. neotesselata for the triploid members).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. scalaris Plateau Spotted Whiptail

Aspidoscelis scalaris (as A. septemvittata) was treated as a subspecies of A. gularis by Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) but as a species by Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist. :27–81). Forstner et al. (1998, J. Herpetol. 32: 418–425) presented evidence of hybridization between A. scalaris and A. gularis. A detailed phylogeographic study of A. scalaris and closely related species is needed.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. scalaris septemvittata Big Bend Spotted Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81). Smith et al. (1996, Herpetol. Rev. 27: 129; priority of the names A. scalaris and A. semifasciata over A. septemvittata and A. sericea and precedence of A. scalaris over A. semifasciata and A. septemvittata over A. sericea). Aspidoscelis scalaris (as A. septemvittata) was treated as a subspecies of A. gularis* by Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) but as a species by Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. sexlineata Six-Lined Racerunner

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. sexlineata sexlineata Eastern Six-Lined Racerunner

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. sexlineata stephensae Texas Yellow-Headed Racerunner

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81), Trauth (1992, Texas J. Sci. 44: 437–443; description of A. sexlineata stephensae),Trauth (1995, Bull. Chicago Herpetol. Soc. 30: 68; spelling of A. sexlineata stephensae).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. sexlineata viridis Prairie Racerunner

See note on A. neomexicana concerning hybridization between that species and A. s. viridis.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. sonorae Sonoran Spotted Whiptail

Unisexual.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. stictogramma Giant Spotted Whiptail

Based on differences in body size, scutellation, and color patterns, Walker and Cordes (2011, Herp. Rev. 42: 33–39) inferred that A. stictogramma (formerly A. burti stictogramma) is a separate species from A. burti.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. tesselata Common Checkered Whiptail

Unisexual. Aspidoscelis dixoni was recognized as a species by Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81) and Walker et al. (1994, Texas J. Sci. 46: 27–33) because its origin was thought to have resulted from a separate hybridization event than the one involved in the origin of the clone represented by the type of A. tesselata. However, Cordes and Walker (2006, Copeia 2006: 14–26) presented evidence in the form of skin-graft histocompatibility that A. dixoni and A. tesselata resulted from a single hybridization event. We have therefore treated the name A. dixoni as a synonym of A. tesselata following Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60). Taylor et al. (2001, Am. Mus. Novit. 3345: 1–65) presented evidence for hybridization between A. tesselata and A. marmorata, but there is no indication that this hybridization has produced a new hybrid species. Cole et al. (2007, Am. Mus. Novit. 3555: 1–31) presented evidence for hybridization between A. tesselata (one of the pattern classes formerly recognized as A. dixoni) and A. tigris punctilinealis and hypothesized that it may be negatively impacting the former taxon.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. tigris Tiger Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81). Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of A. xanthonota as a separate species from A. burti).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. tigris munda California Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81); proposal of A. t. munda as a replacement name for the invalid name A. (t.) undulata Hallowell 1854).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. tigris punctilinealis Sonoran Tiger Whiptail

See note on A. tesselata concerning hybridization between that species and A. t. punctilinealis.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. tigris septentrionalis Plateau Tiger Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. tigris stejnegeri San Diegan Tiger Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81), Maslin and Walker (1981, Am. Midl. Nat. 105: 84–92; treatment of A. t. stejnegeri as the name of the subspecies of A. tigris occurring in coastal southern California).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. tigris tigris Great Basin Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81), Taylor and Walker (1996, Copeia 1996: 140–148; synonymy of A. t. gracilis with A. t. tigris.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

a. uniparens Desert Grassland Whiptail

Unisexual.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. velox Plateau Striped Whiptail

Unisexual. Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) treated the name Aspidoscelis (sackii) innotata as a synonym of A. velox, but Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81) applied the name A. velox to populations of triploid parthenogens and treated A. innotata as the name of a separate diploid species. Cuellar (1977, Evolution 31: 24–31) found histoincompatibility (rejection of skin grafts) between A. velox-like lizards from Colorado, New Mexico, and Utah, which Cuellar and Wright (1992, C. R. Soc. Biogeogr. 68: 157–160) interpreted as potential evidence for different ploidy levels. The type locality of A. velox is in Arizona, while that of A. innotata is in Utah, and lizards from New Mexico are known to be triploid (Neaves, 1969, J. Exper. Zool. 171: 175–184; Dessauer and Cole, 1989, in R. M. Dawley and J. P. Bogart [eds.], Evolution and Ecology of Unisexual Vertebrates, New York State Museum, Pp. 49–71). If lizards from the type locality of A. innotata turn out to be diploid, it would be reasonable to recognize a separate diploid species and apply the name A. innotata (Plateau Unspotted Whiptail) to it.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

a. xanthonota Red-Backed Whiptail

Taxonomy for Aspidoscelis follows Maslin and Secoy (1986, Contrib. Zool. Univ. Colorado Mus. 1: 1–60) and Wright (1993, in J. W. Wright and L. J. Vitt [eds.], Biology of Whiptail Lizards [Genus Cnemidophorus], Oklahoma Mus. Nat. Hist., Pp. 27–81).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

b. vittatus Brown Basilisk

Alien Species:

The Brown Basilisk is native to Central and northern South America and is established in Florida.

Fred Kraus, 2015-01-19

c. draconoides Zebra-Tailed Lizard

Recent molecular phylogeographic studies shed some preliminary light on the relationships and status of the three U.S. subspecies of C. draconoides. Based on mtDNA, Lindell et al. (2005, Mol. Phylogenet. Evol. 36: 682–694) found that both C. d. myurus and C. d. ventralis are nested within C. d. rhodostictus, C. d. ventralis deeply so; however, both C. d. myurus and C. d. ventralis were represented by small samples, and there were large geographic gaps between these samples and those representing C. d. rhodostictus. Blaine (2008, Ph.D. dissertation, Washington Univ.) found that samples representing C. d. rhodostictus and C. d. myurus formed a mtDNA haplotype clade, as did those representing C. d. ventralis, but he had few samples from Baja California and none from the Mexican mainland. Based on genome-wide SNP data, Gottscho (2015, Ph.D. dissertation, Univ. California, Riverside and San Diego State Univ.) inferred that C. d. rhodostictus is distinct from more southerly subspecies endemic to Baja California.

Note on genus: Taxonomy for Callisaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. draconoides myurus Northern Zebra-Tailed Lizard

Recent molecular phylogeographic studies shed some preliminary light on the relationships and status of the three U.S. subspecies of C. draconoides. Based on mtDNA, Lindell et al. (2005, Mol. Phylogenet. Evol. 36: 682–694) found that both C. d. myurus and C. d. ventralis are nested within C. d. rhodostictus, C. d. ventralis deeply so; however, both C. d. myurus and C. d. ventralis were represented by small samples, and there were large geographic gaps between these samples and those representing C. d. rhodostictus. Blaine (2008, Ph.D. dissertation, Washington Univ.) found that samples representing C. d. rhodostictus and C. d. myurus formed a mtDNA haplotype clade, as did those representing C. d. ventralis, but he had few samples from Baja California and none from the Mexican mainland. Based on genome-wide SNP data, Gottscho (2015, Ph.D. dissertation, Univ. California, Riverside and San Diego State Univ.) inferred that C. d. rhodostictus is distinct from more southerly subspecies endemic to Baja California.

Note on genus: Taxonomy for Callisaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. draconoides rhodostictus Western Zebra-Tailed Lizard

Recent molecular phylogeographic studies shed some preliminary light on the relationships and status of the three U.S. subspecies of C. draconoides. Based on mtDNA, Lindell et al. (2005, Mol. Phylogenet. Evol. 36: 682–694) found that both C. d. myurus and C. d. ventralis are nested within C. d. rhodostictus, C. d. ventralis deeply so; however, both C. d. myurus and C. d. ventralis were represented by small samples, and there were large geographic gaps between these samples and those representing C. d. rhodostictus. Blaine (2008, Ph.D. dissertation, Washington Univ.) found that samples representing C. d. rhodostictus and C. d. myurus formed a mtDNA haplotype clade, as did those representing C. d. ventralis, but he had few samples from Baja California and none from the Mexican mainland. Based on genome-wide SNP data, Gottscho (2015, Ph.D. dissertation, Univ. California, Riverside and San Diego State Univ.) inferred that C. d. rhodostictus is distinct from more southerly subspecies endemic to Baja California.

Note on genus: Taxonomy for Callisaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. draconoides ventralis Eastern Zebra-Tailed Lizard

Recent molecular phylogeographic studies shed some preliminary light on the relationships and status of the three U.S. subspecies of C. draconoides. Based on mtDNA, Lindell et al. (2005, Mol. Phylogenet. Evol. 36: 682–694) found that both C. d. myurus and C. d. ventralis are nested within C. d. rhodostictus, C. d. ventralis deeply so; however, both C. d. myurus and C. d. ventralis were represented by small samples, and there were large geographic gaps between these samples and those representing C. d. rhodostictus. Blaine (2008, Ph.D. dissertation, Washington Univ.) found that samples representing C. d. rhodostictus and C. d. myurus formed a mtDNA haplotype clade, as did those representing C. d. ventralis, but he had few samples from Baja California and none from the Mexican mainland. Based on genome-wide SNP data, Gottscho (2015, Ph.D. dissertation, Univ. California, Riverside and San Diego State Univ.) inferred that C. d. rhodostictus is distinct from more southerly subspecies endemic to Baja California.

Note on genus: Taxonomy for Callisaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. versicolor Variable Bloodsucker

The Variable Bloodsucker is native to southern and southeastern Asia and is established in Florida. The specific epithet is in quotation marks because Zug et al. (2006, Proc. Cal. Acad. Sci. 57: 35–68) demonstrated that C. “versicolor” is a complex of several species. The introduced population has yet to be identified in light of this new information.

Note on genus: The English name is derived from the brilliant orange or crimson colors that breeding males develop around the head and shoulders.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-01-13

c. ocellatus Ocellated Skink

Alien Species:

The Ocellated Skink is native to the Mediterranean region, Middle East, and northern Africa and is established in Arizona and Florida.

Fred Kraus, 2015-01-19

c. calyptratus Veiled Chameleon

Alien Species:
The Veiled Chameleon is native to the southwestern Arabian Peninsula and is established in Florida and Hawaii.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

c. lemniscatus Rainbow Whiptail

Alien Species: The Rainbow Whiptail is native to South America and is established in Florida. Several species, both uni- and bisexual, have been described for different parts of the taxon that was formerly known as “C.” lemniscatus (Cole and Dessauer, 1993, Am. Mus. Novit. 3081: 1–30; Markezich et al., 1997, Am. Mus. Novit. 3207: 1–60), and the introduced population is bisexual but has not yet been associated with one or more of those species.

Note on genus: Taxonomy for “Cnemidophorus” follows Peters and Donoso-Barros (1970, Bull. United States Natl. Mus. 297(Part II): 1–293). Reeder et al. (2002, Am. Mus. Novit. 3365: 1–61) presented evidence that Cnemidophorus, even after the removal of Aspidoscelis, is not monophyletic, although they did not propose a taxonomic change to rectify this situation. We have placed the name “Cnemidophorus” in quotation marks to indicate the apparently non-monophyletic status of the taxon.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

c. brevis Texas Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press: 369–469) with modifications described.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. reticulatus Reticulate Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press: 369–469) with modifications described.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. switaki Switak's Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press: 369–469) with modifications described.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. switaki switaki Peninsula Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press: 369–469) with modifications described.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. variegatus Western Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press: 369–469) with modifications described. Leavitt (2015, Ph.D. dissertation, Univ. California, Davis and San Diego State Univ.) presented evidence, based on mt and nuDNA sequences, that C. v. variegatus and C. v. abbotti constitute lineages with limited bi-directional nuclear gene flow and that C. v. bogerti and C. v. utahensis are not differentiated from C. v. variegatus.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. variegatus abbotti San Diego Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press, Pp. 369–469).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

c. variegatus variegatus Desert Banded Gecko

Taxonomy for Coleonyx follows Grismer (1988, in Phylogenetic Relationships of the Lizard Families, R. Estes and G. Pregill [eds.], Stanford Univ. Press, Pp. 369–469).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

c. texanus Greater Earless Lizard

Taxonomy for Cophosaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley). Blaine (2008, Ph.D. dissertation, Washington Univ.) found that most C. texanus sampled within the United States formed three non-overlapping mtDNA haplotype clades, the relationships among which were poorly supported. If the central clade is more closely related to the western clade, then the two primary clades would correspond roughly with the two subspecies of C. texanus that occur in the United States. Samples from the vicinity of Eagle Pass, Maverick County, Texas, formed a separate, earlier diverging clade that could represent a separate species or subspecies.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. texanus scitulus Chihuahuan Greater Earless Lizard

Taxonomy for Cophosaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley). Blaine (2008, Ph.D. dissertation, Washington Univ.) found that most C. texanus sampled within the United States formed three non-overlapping mtDNA haplotype clades, the relationships among which were poorly supported. If the central clade is more closely related to the western clade, then the two primary clades would correspond roughly with the two subspecies of C. texanus that occur in the United States. Samples from the vicinity of Eagle Pass, Maverick County, Texas, formed a separate, earlier diverging clade that could represent a separate species or subspecies.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

c. texanus texanus Texas Greater Earless Lizard

Taxonomy for Cophosaurus follows de Queiroz (1989, Ph.D. dissertation, Univ. California, Berkeley). Blaine (2008, Ph.D. dissertation, Washington Univ.) found that most C. texanus sampled within the United States formed three non-overlapping mtDNA haplotype clades, the relationships among which were poorly supported. If the central clade is more closely related to the western clade, then the two primary clades would correspond roughly with the two subspecies of C. texanus that occur in the United States. Samples from the vicinity of Eagle Pass, Maverick County, Texas, formed a separate, earlier diverging clade that could represent a separate species or subspecies.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

c. bicinctores Great Basin Collared Lizard

Taxonomy for Crotaphytus follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143); for precedence of C. vestigium over C. fasciolatus see McGuire (2000, Bull. Zool. Nomencl. 57: 158–161) and ICZN (2002, Bull. Zool. Nomencl. 59: 228–229). McGuire et al. (2007, Evolution 61: 2879–2897) interpreted incongruences between their mtDNA phylogeny and currently recognized species boundaries in Crotaphytus as evidence for introgression of C. collaris haplotypes into both C. reticulatus and C. bicinctores resulting from past hybridization during glacial maxima.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. collaris Eastern Collared Lizard

Taxonomy for Crotaphytus follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143); for precedence of C. vestigium over C. fasciolatus see McGuire (2000, Bull. Zool. Nomencl. 57: 158–161) and ICZN (2002, Bull. Zool. Nomencl. 59: 228–229). McGuire et al. (2007, Evolution 61: 2879–2897) interpreted incongruences between their mtDNA phylogeny and currently recognized species boundaries in Crotaphytus as evidence for introgression of C. collaris haplotypes into both C. reticulatus and C. bicinctores resulting from past hybridization during glacial maxima.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. nebrius Sonoran Collared Lizard

Taxonomy for Crotaphytus follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143); for precedence of C. vestigium over C. fasciolatus see McGuire (2000, Bull. Zool. Nomencl. 57: 158–161) and ICZN (2002, Bull. Zool. Nomencl. 59: 228–229). McGuire et al. (2007, Evolution 61: 2879–2897) interpreted incongruences between their mtDNA phylogeny and currently recognized species boundaries in Crotaphytus as evidence for introgression of C. collaris haplotypes into both C. reticulatus and C. bicinctores resulting from past hybridization during glacial maxima.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. reticulatus Reticulate Collared Lizard

Taxonomy for Crotaphytus follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143). McGuire et al. (2007, Evolution 61: 2879–2897) interpreted incongruencies between their mtDNA phylogeny and currently recognized species boundaries in Crotaphytus as evidence for introgression of C. collaris haplotypes into both C. reticulatus and C. bicinctores resulting from past hybridization during glacial maxima.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. vestigium Baja California Collared Lizard

Taxonomy for Crotaphytus follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143); for precedence of C. vestigium over C. fasciolatus see McGuire (2000, Bull. Zool. Nomencl. 57: 158–161) and ICZN (2002, Bull. Zool. Nomencl. 59: 228–229). McGuire et al. (2007, Evolution 61: 2879–2897) interpreted incongruences between their mtDNA phylogeny and currently recognized species boundaries in Crotaphytus as evidence for introgression of C. collaris haplotypes into both C. reticulatus and C. bicinctores resulting from past hybridization during glacial maxima.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

c. poecilopleurus Pacific Snake-Eyed Skink

Alien Species:

The Pacific Snake-eyed Skink is native to many Pacific islands and is established in Hawaii.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

c. conspicuosa Isla San Esteban Spiny-Tailed Iguana

Alien Species:

A population of Ctenosaura established at the Arizona-Sonora Desert Museum in Arizona contains mitochondrial DNA from the Isla San Esteban Spiny-tailed Iguana, but it remains uncertain whether this represents a pure population of this species or a hybrid swarm with the C. macrolopha (Edwards et al., 2005, Sonoran Herpetologist 18: 122–125). Both are often considered subspecies of C. hemilopha.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-14

c. macrolopha Sonoran Spiny-Tailed Iguana

Alien Species:

A population of Ctenosaura established at the Arizona-Sonora Desert Museum in Arizona contains mitochondrial DNA from the Sonoran Spiny-tailed Iguana, but it remains uncertain whether this represents a pure population of this species or a hybrid swarm with the C. conspicuosa (Edwards et al., 2005, Sonoran Herpetologist 18: 122–125). Both are often considered subspecies of C. hemilopha.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

c. pectinata Mexican Spiny-Tailed Iguana

Alien Species:

The Mexican Spiny-tailed Iguana is native to Central America and is established in Florida and Texas.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-14

c. similis Gray's Spiny-tailed Iguana

Alien Species:

Gray’s Spiny-tailed Iguana is native to Central America and is established in Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

c. scabrum Rough-Tailed Gecko

Alien Species:

The Rough-tailed Gecko is native to the Middle East and northeastern Africa and is established in Arizona, Nevada, and Texas.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-14

d. dorsalis Desert Iguana

Taxonomy for Dipsosaurus follows de Queiroz (1995, Publ. Espec. Mus. Zool. Univ. Nac. Autón. México 9: 1–48).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

d. dorsalis dorsalis Northern Desert Iguana

Taxonomy for Dipsosaurus follows de Queiroz (1995, Publ. Espec. Mus. Zool. Univ. Nac. Autón. México 9: 1–48).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. coerulea Northern Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. coerulea coerulea San Francisco Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. coerulea palmeri Sierra Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. coerulea principis Northwestern Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. coerulea shastensis Shasta Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. kingii Madrean Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. kingii nobilis Arizona Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. multicarinata Southern Alligator Lizard

A study of mtDNA sequences by Feldman and Spicer (2006, Mol. Ecol. 15: 2201– 2222) and one of mt and nDNA sequences by Leavitt (2015, Ph.D. dissertation, Univ. California, Davis and San Diego State Univ.) failed to support previously recognized subspecies boundaries within E. multicarinata (Fitch, 1938, Am. Midl. Nat. 20: 381– 424). Alleles of specimens from the central Coast Ranges of California (formerly E. m. multicarinata) are more closely related to those of specimens from southern (E. m. webbii) rather than northern (E. m. multicarinata) California, while alleles of specimens from the Sierra Nevada (formerly E. m. webbii) are more closely related to those of specimens from northern (E. m. multicarinata) rather than southern (E. m. webbii) California. In addition, alleles representing E. m. scincicauda are phylogenetically intermixed with, as well as nested within, those of E. m. multicarinata. For these reasons, we have eliminated E. m. scincicauda and have changed the standard English names of the recognized subspecies. There is a potential problem with the scientific names of the subspecies: the type locality of E. m. multicarinata, given only as “Californie”, was thought likely by Fitch (op. cit.) to be in the vicinity of Monterey, which is within the distribution of E. m. webbii as currently recognized.

Kevin de Quieroz (Chair), Tod W. Reeder and Adam D. Leaché, 2018-01-13

e. multicarinata multicarinata Forest Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. multicarinata webbii Woodland Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

e. panamintina Panamint Alligator Lizard

Taxonomy for Elgaria follows Good (1988, Univ. California Pub. Zool. 121: 1–139). The results of Feldman and Spicer (2006, Mol. Ecol. 15: 2201–2222) indicate that E. panamintina is derived from within E. multicarinata.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

e. cyanura Copper-Tailed Skink

Alien Species:

The Copper-tailed Skink is native to the Pacific islands, was established in Hawaii, and may now be extinct there (Fisher and Ineich, 2012, Oryx 46: 187–195). Taxonomy for Emoia cyanura and E. impar follows Ineich and Zug (1991, Copeia 1991: 1132–1136).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

e. impar Azure-Tailed Skink

Alien Species:

The Azure-tailed Skink is native to the Pacific islands and is established in Hawaii.Taxonomy for Emoia cyanura and E. impar follows Ineich and Zug (1991, Copeia 1991: 1132–1136).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

f. oustaleti Oustalet's Chameleon

Alien species: Oustalet’s Chameleon is native to Madagascar and is established in Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-01-13

f. pardalis Panther Chameleon

Alien species: The Panther Chameleon is native to Madagascar and is established in Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-01-13

g. copeii Cope's Leopard Lizard

Taxonomy for Gambelia follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143) with modifications by Frost and Collins (1988, Herpetol. Rev. 19: 73–74; spelling of the specific epithet of G. sila). McGuire et al. (2007 Evolution 61: 2879–2897) found the mtDNA of G. copeii to be deeply nested within that of G. wislizenii and suggested that perhaps the former should not be recognized as a separate species. A study of gene flow (or the absence thereof) between the two forms would clarify the situation.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

g. sila Blunt-Nosed Leopard Lizard

Grimes et al. (2014, Southwestern Nat. 59: 38–46) found that the mtDNA of this species forms two non-overlapping haplotype clades. Taxonomy for Gambelia follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143) with modifications by Frost and Collins (1988, Herpetol. Rev. 19: 73–74; spelling of the specific epithet of G. sila).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

g. wislizenii Long-Nosed Leopard Lizard

Taxonomy for Gambelia follows McGuire (1996, Bull. Carnegie Mus. Nat. Hist. 32: 1–143)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

g. mutilata Mutilating Gecko

Alien Species:
The Mutilating Gecko is native from South Asia through the Pacific islands, has been reported from three states, and is established in Hawaii. The date of publication of the name Hemidactylus mutilatus (=Gehyra mutilata) is sometimes given as 1835 (e.g., Kluge, 1991, Smithsonian Herpetol. Info. Serv. 85: 1–35) presumably based on the idea that the species was first described by Wiegmann in Nova Acta Acad. Caes. Leop. Carol. Nat. Cur. the date of which is either 1834 or 1835; however, the first valid use of the name is in Wiegmann (1834, Herpetologica Mexicana; see Bauer and Adler, 2001, Arch. Nat. Hist., 28: 313–326 for a discussion of the dates of the relevant publications).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

g. badenii Golden Gecko

Alien Species:

The Golden Gecko is native to Vietnam and is established in Florida.

Fred Kraus, 2015-01-19

g. gecko Tokay Gecko

Alien Species:

The Tokay Gecko is native to southeastern Asia and has been introduced to Florida and Hawaii. It is established in Florida but the single known incipient population in Hawaii has apparently been eradicated.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-14

g. infernalis Texas Alligator Lizard

Taxonomy for Gerrhonotus follows Good (1994, Herpetol. Monog. 8: 180–202).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

g. albogularis Yellow-Headed Gecko

Alien Species:

The Yellow-headed Gecko is native to Central and South America and the Caribbean and is established in Florida.

Fred Kraus, 2015-01-19

h. suspectum Gila Monster

Douglas et al. (2010, Mol. Phylogenet. Evol. 55: 153–167) stated that they found no mtDNA evidence for the two subspecies of H. suspectum; however, their results are diffcult to evaluate because little information is provided on the collection localities of the sampled specimens. Further study is needed.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

h. suspectum cinctum Banded Gila Monster

Taxonomy for Heloderma follows Bogert and Martín del Campo (1956, Bull. Am. Mus. Nat. Hist. 109: 1–238). Douglas et al. (2010, Mol. Phylogenet. Evol. 55: 153–167) stated that they found no mtDNA evidence for the two recognized subspecies of H. suspectum; however, their results are difficult to evaluate because little information is provided on the collection localities of the sampled specimens. Further study is needed.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. suspectum suspectum Reticulate Gila Monster

Taxonomy for Heloderma follows Bogert and Martín del Campo (1956, Bull. Am. Mus. Nat. Hist. 109: 1–238). Douglas et al. (2010, Mol. Phylogenet. Evol. 55: 153–167) stated that they found no mtDNA evidence for the two recognized subspecies of H. suspectum; however, their results are difficult to evaluate because little information is provided on the collection localities of the sampled specimens. Further study is needed.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. frenatus Common House Gecko

Alien Species:

The Common House Gecko is native to South and Southeast Asia, has been reported from four states, and is established in Florida, Hawaii, and Texas.

Fred Kraus, 2015-01-19

h. garnotii Indo-Pacific House Gecko

Alien Species:

The Indo-Pacific Gecko is native to South and Southeast Asia and is established in California, Florida, Georgia, Hawaii, and Texas.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

h. mabouia Wood Slave

Alien Species:

The Wood Slave is native to Africa (and perhaps parts of South America and the Caribbean, cf. Kluge, 1969, Misc. Publ. Univ. Michigan Mus. Zool. 138: 1–78), has been reported from three states, and is established in Florida and Texas.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

h. parvimaculatus Sri Lankan Spotted House Gecko

Alien Species:

The Sri Lankan Spotted House Gecko is native to Sri Lanka and southern India and is established in Louisiana.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

h. platyurus Asian Flat-Tailed House Gecko

Alien Species:

The Asian Flat-tailed House Gecko is native to Southeast Asia and is established in Florida. This species was recently removed from Cosymbotus by Carranza and Arnold (2006, Mol. Phylog. Evol. 38: 531–545).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

h. turcicus Mediterranean Gecko

Alien Species:

The Mediterranean Gecko is native to the Mediterranean region, has been reported from 24 states, and is established in Alabama, Arizona, Arkansas, California, Florida, Georgia, Illinois, Kansas, Kentucky, Louisiana, Maryland, Mississippi, Missouri, Nevada, New Mexico, North Carolina, Oklahoma, South Carolina, Tennessee, Texas, Utah, and Virginia.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

h. typus Indo-Pacific Tree Gecko

Alien Species:

Unisexual. The Indo-Pacific Tree Gecko is native to Southeast Asia and the Pacific, has been reported from two states, and is established in Hawaii.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-14

h. elegans Elegant Earless Lizard

Blaine (2008, Ph.D. dissertation, Washington Univ.) found large levels of mtDNA sequence divergence between samples of this putative species from Arizona and southern Sonora (H. e. thermophila) versus those from southern Sinaloa (H. e. elegans), though large sampling gaps make it difficult to determine whether these forms represent separate species. His data also support the synonymy of H. m. pulchra with H. e. thermophila).

Note on genus: Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press) with modifications by Duellman (1955, Occ. Pap. Mus. Zool. Univ. Michigan 569: 1–14; synonymy of H. m. pulchra with H. m. thermophila), Axtell (1956, Bull. Chicago Acad. Sci 10: 163–179; proposal of H. maculata perspicua and treatment of H. lacerata as a species), Clarke (1965, Emporia St. Res. Stud. 13: 1–66; removal of H. texana to Cophosaurus), Lowe (1964, in C. H. Lowe [ed.], The Vertebrates of Arizona, Univ. Arizona Press, Tucson, 153–174; recognition of H. elegans as a species; for supporting evidence see Adest, 1978, Ph.D. dissertation, Univ. California, Los Angeles, Wilgenbusch and de Queiroz, 2000, Syst. Biol. 49: 592–612, and Axtell, 1998, Interpretive Atlas of Texas Lizards 18: 1–19), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

h. elegans thermophila Sonoran Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press). Blaine (2008, Ph.D. dissertation, Washington Univ.) found large levels of mtDNA sequence divergence between samples of this putative species from Arizona and southern Sonora (H. e. thermophila) versus those from southern Sinaloa (H. e. elegans), though large sampling gaps make it difficult to determine whether these forms represent separate species. His data also support the synonymy of H. m. pulchra with H. e. thermophila).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. lacerata Spot-Tailed Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press) with modifications by Axtell (1956, Bull. Chicago Acad. Sci 10: 163–179; description of H. maculata perspicua and treatment of H. lacerata as a species)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. lacerata lacerata Northern Spot-Tailed Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press) with modifications by Axtell (1956, Bull. Chicago Acad. Sci 10: 163–179; description of H. maculata perspicua and treatment of H. lacerata as a species)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. lacerata subcaudalis Southern Spot-Tailed Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press) with modifications by Axtell (1956, Bull. Chicago Acad. Sci 10: 163–179; description of H. maculata perspicua and treatment of H. lacerata as a species)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. maculata Common Lesser Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press). Blaine (2008, Ph.D. dissertation, Washington Univ.) found that Holbrookia maculata from the United States formed three non-overlapping mtDNA haplotype clades inhabiting the Great Plains, the northern Chihuahuan Desert, and the southern Colorado Plateau. Because his results contradict the taxonomy previously adopted in this list, we have applied the oldest available names to the three haplotype clades and treated them as subspecies.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

h. maculata campi Plateau Earless Lizard

Blaine (2008, Ph.D. dissertation, Washington Univ.) found that Holbrookia maculata from the United States formed three non-overlapping mtDNA haplotype clades inhabiting the Great Plains, the northern Chihuahuan Desert, and the southern Colorado Plateau. Because his results contradict the taxonomy previously adopted in this list, we have applied the oldest available names to the three haplotype clades and treated them as subspecies.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. maculata flavilenta Chihuahuan Lesser Earless Lizard

Blaine (2008, Ph.D. dissertation, Washington Univ.) found that Holbrookia maculata from the United States formed three non-overlapping mtDNA haplotype clades inhabiting the Great Plains, the northern Chihuahuan Desert, and the southern Colorado Plateau. Because his results contradict the taxonomy previously adopted in this list, we have applied the oldest available names to the three haplotype clades and treated them as subspecies.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. maculata maculata Great Plains Earless Lizard

Blaine (2008, Ph.D. dissertation, Washington Univ.) found that Holbrookia maculata from the United States formed three non-overlapping mtDNA haplotype clades inhabiting the Great Plains, the northern Chihuahuan Desert, and the southern Colorado Plateau. Because his results contradict the taxonomy previously adopted in this list, we have applied the oldest available names to the three haplotype clades and treated them as subspecies.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. maculata perspicua Prairie Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press). Axtell (1956, Bull. Chicago Acad. Sci 10: 163–179; description of H. maculata perspicua and treatment of H. lacerata as a species). This subspecies was not sampled by Blaine (2008, Ph.D. dissertation, Washington Univ.) and is thus presently retained until future studies can address its status.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

h. maculata ruthveni Bleached Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press). Although mtDNA haplotypes of H. m ruthveni are nested within those of the taxon that is here called H. m. flavilenta (Blaine, 2008, Ph.D. dissertation, Washington Univ.), Rosenblum and Harmon (2010, Evolution 65: 946–960) found that earless lizards from the White Sands had diverged both morphologically and genetically from their counterparts on adjacent darker soils and concluded that the populations are well on their way toward completing speciation. On the other hand, data from ecotonal individuals suggest that the populations continue to exchange genes (i.e., that speciation is incomplete), and therefore it seems appropriate to treat the bleached form as a subspecies in the sense of a partially separated lineage.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

h. propinqua Keeled Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

h. propinqua propinqua Northern Keeled Earless Lizard

Taxonomy for Holbrookia follows Smith (1946, Handbook of Lizards, Cornell Univ. Press). Blaine (2008, Ph.D. dissertation, Washington Univ.) found that mtDNA from H. p. propinqua forms two non-overlapping haplotype clades, one from the red sands south of the Balcones Escarpment and another from the white sands near the southeastern part of the Balcones Escarpment south into the Gulf Coastal Plain.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

i. iguana Green Iguana

Alien Species:

The Green Iguana is native to Central and South America, has been reported from six states, and is established in Florida and Hawaii.

Fred Kraus, 2015-01-19

l. bilineata Western Green Lacerta

Alien Species:

The Western Green Lacerta is native to Western Europe, has been reported from two states, and is established in Kansas.

Fred Kraus, 2015-01-19

l. delicata Plague Skink

Alien Species:

The Plague Skink is native to eastern Australia and is established in Hawaii.

Fred Kraus, 2015-01-19

l. carinatus Northern Curly-Tailed Lizard

Alien Species:

The Northern Curly-tailed Lizard is native to Cuba, Bahamas, and the Cayman Islands and is established in Florida.

Fred Kraus, 2015-01-19

l. schreibersii Red-sided Curly-tailed Lizard

Alien Species:

The Red-sided Curly-tailed Lizard is native to Hispaniola and is established in Florida.

Fred Kraus, 2015-01-19

l. belliana Butterfly Lizard

Alien Species:

The Butterfly Lizard is native to Southeast Asia and is established in Florida.

Fred Kraus, 2015-01-19

l. rubritaeniata Red-Banded Butterfly Lizard

Alien Species:

The Red-banded Butterfly Lizard is native to Indochina and is established in Florida.

Fred Kraus, 2018-01-17

l. lugubris Mourning Gecko

Alien Species:

Unisexual. The Mourning Gecko is native from South Asia through much of the Pacific, has been reported from four states, and is established in Florida and Hawaii. This taxon is a unisexual complex of diploid and triploid populations of apparently independent origins (Moritz et al., 1993, Biol. J. Linn. Soc. 48: 113–133; Volobouev, 1994, Biogeographica 70: 14).

Fred Kraus, 2015-01-19

l. noctua Moth Skink

Alien Species:

The Moth Skink is native to some of the Pacific Islands and is established in Hawaii.

Fred Kraus, 2015-01-19

m. multifasciata Brown Mabuya

Alien Species:

The Brown Mabuya is native to South and Southeast Asia and is established in Florida.

Fred Kraus, 2015-01-19

o. attenuatus Slender Glass Lizard

Taxonomy for Ophisaurus follows McConkey (1954, Bull. Florida St. Mus. Biol. Sci. 2: 13–23) with modifications by Palmer (1987, Herpetologica, 43: 415–423; proposal of O. mimicus). Macey et al. (1999, Mol. Phylogenet. Evol. 12: 250–272) presented mtDNA evidence that Ophisaurus, if it includes North American, European, African, and Asian species, is not monophyletic. Although they favored placing all species in Anguis, this action is both nomenclaturally disruptive and makes Anguis redundant with Anguinae; we have therefore adopted their alternative proposal of retaining Ophisaurus for the North American and Southeast Asian species.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

o. attenuatus attenuatus Western Slender Glass Lizard

Taxonomy for Ophisaurus follows McConkey (1954, Bull. Florida St. Mus. Biol. Sci. 2: 13–23). Macey et al. (1999, Mol. Phylogenet. Evol. 12: 250–272) presented mtDNA evidence that Ophisaurus, if it includes North American, European, African, and Asian species, is not monophyletic. Although they favored placing all species in Anguis, this action is both nomenclaturally disruptive and makes Anguis redundant with Anguinae; we have therefore adopted their alternative proposal of retaining Ophisaurus for the North American and Southeast Asian species.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

o. attenuatus longicaudus Eastern Slender Glass Lizard

Taxonomy for Ophisaurus follows McConkey (1954, Bull. Florida St. Mus. Biol. Sci. 2: 13–23). Macey et al. (1999, Mol. Phylogenet. Evol. 12: 250–272) presented mtDNA evidence that Ophisaurus, if it includes North American, European, African, and Asian species, is not monophyletic. Although they favored placing all species in Anguis, this action is both nomenclaturally disruptive and makes Anguis redundant with Anguinae; we have therefore adopted their alternative proposal of retaining Ophisaurus for the North American and Southeast Asian species.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

o. compressus Island Glass Lizard

Taxonomy for Ophisaurus follows McConkey (1954, Bull. Florida St. Mus. Biol. Sci. 2: 13–23) with modifications by Palmer (1987, Herpetologica, 43: 415–423; proposal of O. mimicus). Macey et al. (1999, Mol. Phylogenet. Evol. 12: 250–272) presented mtDNA evidence that Ophisaurus, if it includes North American, European, African, and Asian species, is not monophyletic. Although they favored placing all species in Anguis, this action is both nomenclaturally disruptive and makes Anguis redundant with Anguinae; we have therefore adopted their alternative proposal of retaining Ophisaurus for the North American and Southeast Asian species.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

o. mimicus Mimic Glass Lizard

Taxonomy for Ophisaurus follows McConkey (1954, Bull. Florida St. Mus. Biol. Sci. 2: 13–23) with modifications by Palmer (1987, Herpetologica, 43: 415–423; proposal of O. mimicus). Macey et al. (1999, Mol. Phylogenet. Evol. 12: 250–272) presented mtDNA evidence that Ophisaurus, if it includes North American, European, African, and Asian species, is not monophyletic. Although they favored placing all species in Anguis, this action is both nomenclaturally disruptive and makes Anguis redundant with Anguinae; we have therefore adopted their alternative proposal of retaining Ophisaurus for the North American and Southeast Asian species.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

o. ventralis Eastern Glass Lizard

Taxonomy for Ophisaurus follows McConkey (1954, Bull. Florida St. Mus. Biol. Sci. 2: 13–23) with modifications by Palmer (1987, Herpetologica, 43: 415–423; proposal of O. mimicus). Macey et al. (1999, Mol. Phylogenet. Evol. 12: 250–272) presented mtDNA evidence that Ophisaurus, if it includes North American, European, African, and Asian species, is not monophyletic. Although they favored placing all species in Anguis, this action is both nomenclaturally disruptive and makes Anguis redundant with Anguinae; we have therefore adopted their alternative proposal of retaining Ophisaurus for the North American and Southeast Asian species.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. mearnsi Mearns's Rock Lizard

Taxonomy for Petrosaurus follows Jennings (1990, Cat. Am. Amph. Rept. 494; 1990, Cat. Am. Amph. Rept. 495), with modifications by Grismer (1999, Herpetologica 55: 446–469; treatment of P. mearnsi and P. slevini as separate species).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. grandis Madagascan Giant Day Gecko

Alien Species:

The Madagascar Day Gecko is native to Madagascar and is established in Florida and Hawaii. Formerly referred to P. madagascariensis Gray, 1831 prior to partitioning of that species (Raxworthy et al., 2007, Syst. Biol. 56: 907–923).

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

p. guimbeaui Orange-Spotted Day Gecko

Alien Species:

The Orange-spotted Day Gecko is native to Mauritius and is established in Hawaii.

Fred Kraus, 2015-01-19

p. laticauda Gold Dust Day Gecko

Alien Species:

The Gold Dust Day Gecko is native to Madagascar and the Seychelles and is established in Florida and Hawaii.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

p. blainvillii Blainville's Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. cornutum Texas Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. douglasii Pygmy Short-Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii and implied treatment of P. d. brevirostre, P. d. ornatissum, and P. d.ornatum as synonyms of P. hernandesi), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-15

p. goodei Goode's Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. hernandesi Greater Short-Horned Lizard

In a recent revision of the Short-horned Lizards based on morphology, Montanucci (2015, Zootaxa, 4015: 1–177) recognized five species for lizards previously assigned to the single species P. hernandesi (Zamudio et al., 1997, Syst. Biol. 46: 284–305). His taxonomy, however, exhibits both conflicts with inferred phylogenetic relationships based on mt and nDNA (Zamudio et al., op. cit.; Leaché et al., 2015, Syst. Biol. 64: 1032–1047) and internal inconsistencies. For example, the deepest phylogenetic divergence in mtDNA occurs within his subspecies P. h. hernandesi, an extensive putative hybrid zone occurs largely within the distribution of P. h. hernandesi, and the two ostensibly hybridizing species elsewhere exhibit a patchwork distribution, with some populations of P. h. hernandesi completely surrounded by P. o. ornatissimum. In addition, all of the newly recognized species for which molecular data are available are nested within P. hernandesi. For these reasons, all five species recognized by Montanucci (op. cit.) are here treated as subspecies of P. hernandesi. An explicit species delimitation analysis of the Short-horned Lizards based on multilocus genetic data is needed.

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder and Adam D. Leaché, 2018-01-13

p. hernandesi bauri Baur’s Short-horned Lizard

p. hernandesi brevirostris Plains Short-Horned Lizard

p. hernandesi diminutum San Luis Valley Short-Horned Lizard

p. hernandesi hernandesi Hernandez’s Short-horned Lizard

p. hernandesi ornatissimum New Mexico Short-Horned Lizard

p. hernandesi ornatum Great Basin Short-Horned Lizard

p. mcallii Flat-Tailed Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. modestum Round-Tailed Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. platyrhinos Desert Horned Lizard

According to Pianka (1991, Cat. Am. Amph. Rept. 517), the putative diagnostic characters for the subspecies of Phrynosoma platyrhinos are not reliable, which calls the taxa themselves into question. Jezkova et al. (2015, Ecography 38: 1–12) found evidence, based on mtDNA sequences, indicating separate invasions of the Great Basin, with eastern and western populations of P. p. “platyrhinos” more closely related to different populations of P. p. “calidiarum”. Therefore, we have not recognized those subspecies. Phylogenetic analysis of mtDNA sequences by Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; see also Jezkova et al., op. cit.) raised the possibility that lizards from the Yuma Proving Ground represent an unnamed species.

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder and Adam D. Leaché, 2018-01-13

p. solare Regal Horned Lizard

Taxonomy for Phrynosoma follows Reeve (1952, Univ. Kansas Sci. Bull. 34: 817–960) with modifications by Zamudio et al. (1997, Syst. Biol. 46: 284–305; treatment of P. hernandesi as a separate species from P. douglasii; see also Montanucci, 2015, Zootaxa 4015: 1–177), Montanucci (2004, Herpetologica 60: 117–139; treatment of P. blainvillii as a separate species from P. coronatum; see also Leaché et al., 2009, Proc. Natl. Acad. Sci. USA 106: 12418–12423), Mulcahy et al. (2006, Mol. Ecol. 15: 1807–1826; treatment of P. goodei as a separate species from P. platytrhinos), and those described in additional notes below. Leaché and McGuire (2006, Mol. Phylogenet. Evol. 39: 628–644; see also Leaché and Linkem, 2015, Copeia 103: 586–594) named four subclades of Phrynosoma based on the results of phylogenetic analyses of mitochondrial and nuclear genes. We have included names of subclades parenthetically, where applicable.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. nocticolus Peninsula Leaf-Toed Gecko

Taxonomy for Phyllodactylus follows Dixon (1969, Cat. Am. Amph. Rept. 79; 1973, Cat. Am. Amph. Rept. 141) with modifications by Murphy (1983, Occ. Pap. California Acad. Sci. 137: 1–48; treatment of P. nocticolus as a species separate from P. xanti; see also Blair et al., 2009, Zootaxa 2027: 28–42).

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. anthracinus Coal Skink

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. anthracinus anthracinus Northern Coal Skink

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. anthracinus pluvialis Southern Coal Skink

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. callicephalus Mountain Skink

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. egregius Mole Skink

Branch et al. (2003, Conserv. Gen. 4: 199–212) found that the mainland subspecies P. e. lividus, P. e. onocrepsis, and P. e. similis exhibit phylogenetic intermixing of mtDNA haplotypes, suggesting that continued recognition of these taxa may not be warranted. Schrey et al. (2012, J. Herpetol. 46: 241–247) found evidence of genetic differentiation between populations of P. e. lividus north and south of Josephine Creek on the Lake Wales Ridge but did not propose taxonomic recognition of those units.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. egregius egregius Florida Keys Mole Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189). Branch et al. (2003, Conserv. Gen. 4: 199–212) found that the mainland subspecies P. e. lividus, P. e. onocrepsis, and P. e. similis exhibit phylogenetic intermixing of mtDNA haplotypes, suggesting that continued recognition of these taxa may not be warranted. Schrey et al. (2012, Jo. Herpetol. 46: 241–247) found evidence of genetic differentiation between populations of P. e. lividus north and south of Josephine Creek on the Lake Wales Ridge but did not propose taxonomic recognition of those units

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. egregius insularis Cedar Key Mole Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; descriptions of P. egregius lividus and P. e. insularis)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. egregius lividus Blue-Tailed Mole Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; descriptions of P. egregius lividus and P. e. insularis). Schrey et al. (2012, Jo. Herpetol. 46: 241–247) found evidence of genetic differentiation between populations of P. e. lividus north and south of Josephine Creek on the Lake Wales Ridge but did not propose taxonomic recognition of those units.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. egregius onocrepis Peninsula Mole Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. egregius similis Northern Mole Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; description of P. egregius similis)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. fasciatus Common Five-Lined Skink

Howes et al. (2006, Mol. Phylogenet. Evol. 40: 183–194) and Richmond (2006, Evol. Dev. 8: 477–490) presented mitochondrial and nuclear DNA evidence of substantial phylogeographic structure within P. fasciatus. Although neither set of authors drew any taxonomic conclusions from their results, those results suggest the possibility of one or more cryptic species; in particular, samples from the eastern Carolinas are highly divergent in both mtDNA and microsatellites from nearby populations.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. gilberti Gilbert's Skink

Richmond and Reeder (2002, Evolution 56: 1498–1513) presented mtDNA evidence that populations previously referred to Plestiodon gilberti represent three lineages that separately evolved large body size and the loss of stripes in late ontogenetic stages. Although they considered those three lineages to merit species recognition, they did not propose specific taxonomic changes, and subsequently Richmond and Jockusch (2007, Proc. Roy. Soc. Lond. B 274: 1701–1708) and Richmond et al. (2011, Am. Nat. 178: 320–332) have treated them as a single species based on extensive introgressive hybridization between two of the forms and the lack of prezygotic isolation between members of all pairs of them. The results of Richmond and Reeder (2002, op. cit.) contradict the recognition of P. g. arizonensis, which is not differentiated from P. g. rubricaudatus and therefore has been eliminated from this list, and indicate the existence of an unnamed and at least partially separate lineage within P. g. rubricaudatus (their Inyo clade).

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. gilberti cancellosus Variegated Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189)Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; description of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. gilberti gilberti Greater Brown Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; description of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti). Richmond and Reeder (2002, Evolution 56: 1498–1513) presented mitochondrial DNA evidence that populations previously referred to Plestiodon gilberti represent three lineages that separately evolved large body size and the loss of stripes in late ontogenetic stages. Although they considered those three lineages to merit species recognition, they did not propose specific taxonomic changes, and subsequently Richmond and Jockusch (2007, Proc. Roy. Soc. Lond. B 274: 1701–1708) and Richmond et al. (2011, Am. Nat. 178: 320–332) have treated them as a single species based on extensive introgressive hybridization between two of the forms and the lack of prezygotic isolation between members of all pairs of them.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. gilberti placerensis Northern Brown Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; description of P. gilberti placerensis)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. gilberti rubricaudatus Western Red-Tailed Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189),

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. inexpectatus Southeastern Five-Lined Skink

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-02

p. laticeps Broad-Headed Skink

Richmond (2006, Evol. Dev. 8: 477–490) found a substantial division between mtDNA haplotypes of eastern and western P. laticeps but did not draw any taxonomic conclusion from it.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. multivirgatus Many-Lined Skink

Hammerson (1999, Amphibians and Reptiles in Colorado, Univ. Press of Colorado, Niwot) argued, based on diagnosability and the apparent absence of intergrades, that Plestiodon multivirgatus epipleurotus (under the name P. gaigeae) is a different species than P. m. multivirgatus. We have refrained from adopting this proposal pending an explicit analysis.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. multivirgatus epipleurotus Variable Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Hammerson (1999, Amphibians and Reptiles in Colorado, Univ. Press of Colorado) argued, based on diagnosability and the apparent absence of intergrades, that Plestiodon multivirgatus epipleurotus (under the name P. gaigeae) is a different species than P. m. multivirgatus. We have refrained from adopting this proposal pending an explicit analysis.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. multivirgatus multivirgatus Northern Many-Lined Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. obsoletus Great Plains Skink

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. reynoldsi Florida Sand Skink

Branch et al. (2003, Conserv. Gen. 4: 199–212) and Richmond et al. (2009, Conserv. Gen. 10: 1281–1297) found strong phylogeographic structuring in P. reynoldsi, with separate mtDNA clades occupying the Mt. Dora Ridge and the northern, central, and southern portions of the Lake Wales Ridge, but they did not propose to recognize those units taxonomically.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. septentrionalis Prairie Skink

Plestiodon septentrionalis septentrionalis and P. s. obtusirostris have sometimes been recognized as species based on allopatry and morphological diagnosability (e.g., Collins, 1991, Herpetol. Rev. 22: 42–43; 1993, Univ. Kansas Mus. Nat. Hist. Public Edu. Ser. No. 13). Fuerst and Austin (2004, J. Herpetol. 38: 257–268) presented mtDNA evidence of 6–7% sequence divergence between P. s. septentrionalis and P. s. obtusirostris; however, their geographic sampling was inadequate to address genetic continuity versus discontinuity between these taxa. In addition, the name P. s. pallidus, absent from the literature of the last 40 years, apparently has never been explicitly treated as a synonym of either P. s. septentrionalis or P. s. obtusirostris. We have retained the older arrangement of a single species with three subspecies until a rearrangement is proposed based on a study of all three taxa and thorough geographic sampling.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. septentrionalis obtusirostris Southern Prairie Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Plestiodon septentrionalis septentrionalis and P. s. obtusirostris have sometimes been recognized as species based on allopatry and morphological diagnosability (e.g., Collins, 1991, Herpetol. Rev. 22: 42–43; 1993, Univ. Kansas Mus. Nat. Hist. Public Edu. Ser. No. 13). Fuerst and Austin (2004, J. Herpetol. 38: 257–268) presented mtDNA evidence of 6–7% sequence divergence between P. s. septentrionalis and P. s. obtusirostris; however, their geographic sampling was inadequate to address genetic continuity versus discontinuity between these taxa. In addition, the name P. s. pallidus, absent from the literature of the last 40 years, apparently has never been explicitly treated as a synonym of either P. s. septentrionalis or P. s. obtusirostris.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. septentrionalis pallidus Pallid Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Plestiodon septentrionalis septentrionalis and P. s. obtusirostris have sometimes been recognized as species based on allopatry and morphological diagnosability (e.g., Collins, 1991, Herpetol. Rev. 22: 42–43; 1993, Univ. Kansas Mus. Nat. Hist. Public Edu. Ser. No. 13). Fuerst and Austin (2004, J. Herpetol. 38: 257–268) presented mtDNA evidence of 6–7% sequence divergence between P. s. septentrionalis and P. s. obtusirostris; however, their geographic sampling was inadequate to address genetic continuity versus discontinuity between these taxa. In addition, the name P. s. pallidus, absent from the literature of the last 40 years, apparently has never been explicitly treated as a synonym of either P. s. septentrionalis or P. s. obtusirostris.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. septentrionalis septentrionalis Northern Prairie Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Plestiodon septentrionalis septentrionalis and P. s. obtusirostris have sometimes been recognized as species based on allopatry and morphological diagnosability (e.g., Collins, 1991, Herpetol. Rev. 22: 42–43; 1993, Univ. Kansas Mus. Nat. Hist. Public Edu. Ser. No. 13). Fuerst and Austin (2004, J. Herpetol. 38: 257–268) presented mtDNA evidence of 6–7% sequence divergence between P. s. septentrionalis and P. s. obtusirostris; however, their geographic sampling was inadequate to address genetic continuity versus discontinuity between these taxa. In addition, the name P. s. pallidus, absent from the literature of the last 40 years, apparently has never been explicitly treated as a synonym of either P. s. septentrionalis or P. s. obtusirostris.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. skiltonianus Western Skink

Richmond and Reeder (2002, Evolution 56: 1498–1513) presented mitochondrial DNA evidence that P. s. skiltonianus is paraphyletic with respect to both P. s. interparietalis and P. s. utahensis as well as to the species P. lagunensis (Baja California) and to two of the three lineages of P. gilberti.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. skiltonianus interparietalis Coronado Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Richmond and Reeder (2002, Evolution 56: 1498–1513) presented mitochondrial DNA evidence that P. s. skiltonianus is paraphyletic with respect to both P. s. interparietalis and P. s. utahensis as well as to the species P. lagunensis (Baja California) and to two of the three lineages of P. gilberti.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. skiltonianus skiltonianus Skilton's Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189). Richmond and Reeder (2002, Evolution 56: 1498–1513) presented mitochondrial DNA evidence that P. s. skiltonianus is paraphyletic with respect to both P. s. interparietalis and P. s. utahensis as well as to the species P. lagunensis (Baja California) and to two of the three lineages of P. gilberti.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. skiltonianus utahensis Great Basin Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Richmond and Reeder (2002, Evolution 56: 1498–1513) presented mitochondrial DNA evidence that P. s. skiltonianus is paraphyletic with respect to both P. s. interparietalis and P. s. utahensis as well as to the species P. lagunensis (Baja California) and to two of the three lineages of P. gilberti.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. tetragrammus Four-Lined Skink

Moseley et al. (2015, Biol. J. Linn. Soc. 116: 819–833) corroborated the distinction between P. t. brevilineatus and P. t. tetragrammus based on phylogenetic analyses of mtDNA and combined mt and nuDNA.

Notes on genus: Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189) with modifications by Rodgers (1944, Copeia 1944: 101–104; proposal of P. gilberti placerensis), Smith (1946, Univ. Kansas Pub. Mus. Nat. Hist. 1: 85–89; resurrection of P. anthracinus pluvialis), Rodgers and Fitch (1947, Univ. California Pub. Zool. 48: 169–220; proposal of P. gilberti cancellosus and treatment of P. skiltonianus brevipes as a synonym of P. gilberti gilberti), Smith and Slater (1949, Trans. Kansas Acad. Sci. 52: 438–448; proposal of P. septentrionalis pallidus), McConkey (1957, Bull. Florida St. Mus. (Biol. Sci.) 2: 13–23; proposal of P. egregius similis), Lowe and Shannon (1954, Herpetologica 10: 185–187; proposal of P. gilberti arizonensis), Lowe (1955b, Herpetologica 11: 233–235; treatment of P. gaigeae as a subspecies of P. multivirgatus), Mecham (1957, Copeia 1957: 111–123; treatment of P. taylori as a synonym of P. m. gaigeae), Tanner (1958, Great Basin Nat. 17: 59–94; proposals of P. skiltonianus utahensis and P. s. interparietalis), Axtell (1961, Texas J. Sci. 13: 345–351; see also Axtell and Smith, 2004, Southwest. Nat. 49: 100; priority of P. multivirgatus epipleurotus over P. m. gaigeae), Mount (1965, The Reptiles and Amphibians of Alabama, Auburn Univ. Agric. Exper. Station; proposals of P. egregius lividus and P. e. insularis), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19; treatment of P. brevilineatus and P. tetragrammus as subspecies of a single species), Tanner (1987, Great Basin Nat. 47: 383–421; treatment of P. callicephalus as a separate species from P. tetragrammus), Brandley et al. 2005 (Syst. Biol. 54: 373–390; restriction of Eumeces and resurrection of Plestiodon for a clade containing all of the North American species as well as inclusion of the taxon formerly known as Neoseps reynoldsi), and those described in additional notes.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

p. tetragrammus brevilineatus Short-Lined Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19) treated Plestiodon callicephalus as a subspecies of P. tetragrammus.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. tetragrammus tetragrammus Long-Lined Skink

Taxonomy for Plestiodon (often as Eumeces) follows Taylor (1935, Univ. Kansas Sci. Bull. 23: 1–643) and Brandley et al. (2012, Zool. Jo. Linn. Soc. 165: 163–189), Lieb (1985, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 357: 1–19) treated Plestiodon callicephalus as a subspecies of P. tetragrammus.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

p. muralis Common Wall Lizard

Alien Species:

The Common Wall Lizard is native to Europe, has been reported from four states, and is established in Indiana, Kentucky, Ohio, and British Columbia.

Fred Kraus, 2015-01-19

p. siculus Italian Wall Lizard

Alien Species:

The Italian Wall Lizard is native to Europe, has been reported from seven states, and is established in California, Connecticut, Kansas, Missouri, New Jersey, and New York. It was formerly established in Pennsylvania but is now extinct there.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-02-12

r. floridana Florida Wormlizard

Taxonomy for Rhineura follows Gans (1967, Cat. Am. Amph. Rept. 42; 1967, Cat. Am. Amph. Rept. 43). Mulvaney et al. (2005, J. Herpetol. 39: 118–124) found mtDNA evidence of substantial divergence between northern and southern populations of Rhineura floridana and indicated that these groups of populations may be candidates for recognition as separate species.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. merianae Argentine Giant Tegu

The Argentine Giant Tegu is native to South America and is established in Florida.

Fred Kraus (Chair), Kenneth L. Krysko, 2018-01-13

s. ater Common Chuckwalla

Taxonomy for Sauromalus follows Hollingsworth (1998, Herpetol. Monog. 12: 38–191) and the ICZN (2004, Bull. Zool. Nomencl. 61: 74–75; precedence of the name S. ater over S. obesus). Although all mainland populations of Sauromalus are currently considered to constitute a single species, intergradation or the lack thereof between groups based on mtDNA haplotype clades (Petren and Case, 2002, in T. J. Case, M. L. Cody, and E. Ezcurra [eds.], A New Island Biogeography of the Sea of Cortés, Oxford Univ. Press, Pp. 574–579) deserves further study.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. arenicolus Dunes Sagebrush Lizard

Chan et al. (2009, Conserv. Genet. 10: 131–142) found mtDNA and microsatellite evidence of differentiation of S. arenicolus populations into three genetic clusters that appear to be recently separated and still experiencing gene flow.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. bimaculosus Twin-Spotted Spiny Lizard

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. clarkii Clark's Spiny Lizard

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. clarkii clarkii Sonoran Spiny Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. clarkii vallaris Plateau Spiny Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. consobrinus Prairie Lizard

Leaché and Reeder (2002, Syst. Biol. 51: 44–68) noted that the name S. thayerii Baird and Girard 1852 (type locality: Indianola, Calhoun Co., TX) may turn out to be the correct name of this species. Dixon (2013, Amphibians and Reptiles of Texas, Texas A&M University Press, College Station) included Calhoun County, Texas, which includes the type locality of S. thayerii, within the distribution of S. consobrinus; however, he did not indicate a morphological basis for distinguishing S. consobrinus from S. cowlesi (p. 29), nor did he provide genetic evidence for his distributional inference. An assessment of the relationships of fence lizards from the type locality of S. thayerii is needed. Leaché and Reeder (2002, op. cit.) also noted that populations east of the Mississippi River along the Gulf Coast may represent a separate species.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. cowlesi Southwestern Fence Lizard

Leaché and Reeder (2002, Syst. Biol. 51: 44–68) applied the name S. cowlesi to the populations from roughly the region of the Chihuahuan Desert. Although the name S. cowlesi was originally applied to light colored lizards from the White Sands of New Mexico, Leaché and Reeder (op. cit.) presented evidence that mtDNA haplotypes from White Sands lizards are deeply nested within a clade of haplotypes from geographically proximate darker lizards, and Rosenblum (2006, Am. Nat. 167: 1–15) found both phylogenetic mixing of haplotypes between light and dark forms and evidence of gene flow between them. Rosenblum and Harmon (2010, Evolution 65: 946–960) found that fence lizards from the White Sands exhibited discordant patterns of morphological and genetic dfferientiation from their counterparts on adjacent darker soils and concluded that the populations have made incomplete progress toward speciation. Leaché and Cole (2007, Mol. Ecol. 16: 1035–1054) presented evidence for hybridization between S. cowlesi and S. tristichus.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. cyanogenys Blue Spiny Lizard

Olson (1987, Bull. Maryland Herpetol. Soc. 23: 158–167) treated Sceloporus cyanogenys as a subspecies of S. serrifer based on apparent intergrades between the former species and S. serrifer plioporus. Martínez-Méndez and Méndez de la Cruz (2007, Zootaxa 1609: 53–68) inferred S. serrifer plioporus and S. cyanogenys to form a mtDNA clade; however, that clade was relatively distantly related to S. serrifer serrifer and S. serrifer prezygus haplotypes (see also Wiens et al., 2010, Mol. Phylogenet. Evol. 54: 150–161). Therefore, they synonymized the name S. s. plioporus with S. cyanogenys, retaining S. serrifer for a species that occurs south and east of the Isthmus of Tehuantepec.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. graciosus Common Sagebrush Lizard

Chan et al. (2013, Zootaxa 3664: 312–320) found that the currently recognized subspecies of S. graciosus are incongruent with mitochondrial haplotype clades, which often exhibit relatively deep divergences between geographically proximate samples, and that S. graciosus is paraphyletic relative to S. arenicolus. Although these findings suggest that S. graciosus is in need of taxonomic revision, those authors did not propose any taxonomic changes.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. graciosus gracilis Western Sagebrush Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago). Chan et al. (2013, Zootaxa 3664: 312–320) found that the currently recognized subspecies of S. graciosus are incongruent with mitochondrial haplotype clades, which often exhibit relatively deep divergences between geographically proximate samples, and that S. graciosus is paraphyletic relative to S. arenicolus. Although these findings suggest that S. graciosus is in need of taxonomic revision, those authors did not propose any taxonomic changes.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. graciosus graciosus Northern Sagebrush Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago)

Kevin de Queiroz (Chair) and Tod W. Reeder, 2015-01-26

s. graciosus vandenburgianus Southern Sagebrush Lizard

Censky (1986, Cat. Am. Amph. Rept. 386) treated Sceloporus graciosus vandenburgianus as a subspecies of S. graciosus, but Collins (1991, Herpetol. Rev. 22: 42–43) proposed recognizing this taxon as a species, S. vandenburgianus. Wiens and Reeder (1997, Herpetol. Monog. 11: 1–101) followed Collins’s proposal but noted the morphological similarity and geographic proximity of this taxon to populations of S. graciosus gracilis. Chan et al. (2013, Zootaxa 3664: 312–320) found that S. g. vandenburgianus was mitochondrially distinct from S. g. gracilis and paraphyletic relative to a clade formed by eastern populations of S. g. graciosus and S. arenicolus. We have retained vandenburgianus as a subspecies pending a detailed analysis of geographic variation in S. graciosus.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. grammicus Graphic Spiny Lizard

Lizards currently referred to Sceloporus grammicus form a complex series of chromosome races that likely represent multiple species (Sites, 1983, Evolution 37: 38–53; Arévalo et al., 1991, Herpetol. Monog. 5: 79–115). A detailed phylogeographic study of this species complex is needed.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. grammicus microlepidotus Mesquite Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago). Lizards currently referred to Sceloporus grammicus form a complex series of chromosome races that likely represent multiple species (Sites, 1983, Evolution 37: 38–53; Arévalo et al., 1991, Herpetol. Monog. 5: 79–115). A detailed phylogeographic study of this species complex is sorely needed.

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. jarrovii Yarrow's Spiny Lizard

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. magister Desert Spiny Lizard

Leaché and Mulcahy (2007, Mol. Ecol. 16: 5216–5233) found evidence of asymmetrical gene flow between S. magister and both S. bimaculosus and S. uniformis, with S. magister acting as a genetic “sink”. Because these lineages show evidence of both separation (with divergence) and ongoing asymmetrical gene flow, they can be considered partially separated species. Leaché and Mulcahy (op. cit.) also identified a fourth potentially separate lineage in northeastern Baja California (currently unnamed). Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880) recognized the subspecies S. m. magister and S. m. cephaloflavus because their single sample from the Colorado Plateau (assumed to represent the subspecies S. m. cephaloflavus) was inferred to be the sister group of the samples representing S. m. magister. Leaché and Mulcahy (op. cit.), however, found that specimens from closer to the type locality of S. m. cephaloflavus were part of S. uniformis rather than S. magister; consequently, we have not recognized subspecies within S. magister.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. merriami Canyon Lizard

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 51: 44–68; treatment of S. consobrinus, S. cowelsi, and S. tristichus as separate species from S. undulatus), Schulte et al. (2006, Mol. Phylogenet. Evol. 39: 873–880; treatment of S. bimaculosus and S. uniformis as species separate from S. magister and tranversus as a synonym of uniformis; see Leaché and Mulcahy, 2007, Mol. Ecol. 16: 5216–5233 for clarification of the distributional limits of those species), and those described in additional notes below.

Kevin de Quieroz (Chair), Tod W. Reeder, and Adam D. Leaché, 2018-02-10

s. merriami annulatus Big Bend Canyon Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. merriami longipunctatus Presidio Canyon Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago).

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. merriami merriami Merriam's Canyon Lizard

Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago)

Kevin de Quieroz (Chair) and Tod W. Reeder, 2015-01-26

s. occidentalis Western Fence Lizard

Leaché et al. (2010, Biol. Jo. Linn. Soc. 100: 630–641) presented mtDNA evidence that the previously recognized subspecies S. o. taylori is polyphyletic and represents convergent phenotypic evolution among high elevation populations of S. o. biseriatus.

Notes on genus: Taxonomy for Sceloporus follows Schmidt (1953, A Check List of North American Amphibians and Reptiles, Univ. Chicago Press, Chicago) with modifications by Bell (1954, Herpetologica 10: 31–36; resurrection of S. occidentalis bocourtii and S. o. longipes), Shannon and Urbano (1954, Herpetologica 10: 189–191; proposal of S. clarki vallaris), Phelan and Brattstrom (1955, Herpetologica 11: 1–14; proposals of S. magister uniformis, S. m. bimaculosus, and S. m. transversus), Tanner (1955, Great Basin Nat. 15: 32–34; proposal of S. magister cephaloflavus), Lowe and Norris (1956, Herpetologica 12: 125–127; proposal of S. undulatus cowlesi), Maslin (1956, Herpetologica 12: 291–294; proposal of S. undulatus erythrocheilus), Smith and Chrapliwy (1958, Herpetologica 13: 267–271; proposal of subspecies of S. poinsettii), Cole (1963, Copeia 1963: 413–425; treatment of S. virgatus as a species separate from S. undulatus), Degenhardt and Jones (1972, Herpetologica 28: 212–217; proposal of S. graciosus arenicolous), Olson (1973, Herpetologica 29: 116–127; proposal of S. merriami longipunctatus), Sites and Dixon (1981, J. Herpetol. 15: 59–69; treatment of disparilis as a synonym of microlepidotus), Collins (1991, Herpetol. Rev. 22: 42–43; treatment of S. arenicolus as a species separate from S. graciosus; corroborated by Chan et al., 2013, Zootaxa 3664: 312–320), Smith et al. (1992, Bull. Maryland Herpetol. Soc. 28: 123–149; proposal of S. undulatus tedbrowni and correction of the spelling of the name S. arenicolus), Smith et al. (1996, Bull. Maryland Herpetol. Soc. 32: 70–74; treatment of S. slevini as a species separate from S. scalaris; corroborated by Bryson et al., 2012, Mol. Phylogenet. Evol. 62: 447–457 and Grummer et al., 2014, Syst. Biol. 63: 119–133), Wiens et al. (1999, Evolution 53: 1884–1897; restriction of the name S. jarrovii to one of five inferred species formerly referred to by that name), Leaché and Reeder (2002, Syst. Biol. 5