IGUANAS OF THE WORLD Family Iguanidae Subfamily Iguaninae. Amblyrhynchus Bell [Galápagos Marine Iguanas] - PDF

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IGUANAS OF THE WORLD Family Iguanidae Subfamily Iguaninae Checklist compiled by John B. Iverson and based primarily on Etheridge (1982) and Hollingsworth (2004). Content: Nine living genera and 42 living

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IGUANAS OF THE WORLD Family Iguanidae Subfamily Iguaninae Checklist compiled by John B. Iverson and based primarily on Etheridge (1982) and Hollingsworth (2004). Content: Nine living genera and 42 living species are currently recognized; however, undescribed species are known to exist (e.g., see Zarza et al. 2008; Malone and Davis 2004; Gentile et al. 2009). Comment: This assemblage of relatively large, mostly herbivorous iguanas has long been recognized as being monophyletic (all derived from a single common ancestor). As such, most scientists have classified the group in the subfamily Iguaninae of the family Iguanidae (Schulte et al. 2003; Pough et al. 2004; Vitt and Caldwell 2009), although some recent authors (e.g., Frost et al. 2001; Hollingsworth 1998, 2004; Conrad 2008) have elevated the iguanas to full family status. Distribution: New World tropics and subtropics including the West Indies and Galápagos Islands, and the Fiji-Tonga Islands. Comment: Considerable controversy surrounds the phylogenetic relationships among the genera in this family (e.g., see Hollingsworth 2004). Amblyrhynchus Bell [Galápagos Marine Iguanas] Amblyrhynchus Bell 1825, Zool. Jr., London 2:206. Type species (by monotype): Amblyrhynchus cristatus Bell Distribution: Galápagos Islands. Amblyrhynchus cristatus Bell [Galápagos Marine Iguana] Original name: Amblyrhynchus cristatus Bell 1825, Zool. Journal, London 2:206. Holotype: OUM 6176 (Etheridge 1982). Type locality: Mexico. Corrected type locality: Narborough (Fernandina) (Eibl-Eibesfeldt 1956). Distribution: Galápagos Archipelago. Comment: Most authors have not used subspecific epithets, although they have not been formally synonymized (see Rassmann et al. 1997). Amblyrhynchus cristatus cristatus Bell [Fernandina Marine Iguana] Original name: Amblyrhynchus cristatus cristatus Bell. See species account. Distribution: Narborough (Fernandina) Island, Galápagos Archipelago. Amblyrhynchus cristatus albermarlensis Eibl-Eibesfeldt [Isabella Marine Iguana] Original name: Amblyrhynchus cristatus albermarlensis Eibl-Eibesfeldt 1962, Senckenberg. Biol., Frankfurt a. Main 43(3):184. Holotype: Eibl-Eibesfeldt private coll. (Etheridge 1982). Type locality: Insel Albemarle (Isabella). Distribution: Albemarle (Isabella) Island, Galápagos Archipelago. Amblyrhynchus cristatus hassi Eibl-Eibesfeldt [Santa Cruz Marine Iguana] Original name: Amblyrhynchus cristatus hassi Eibl-Eibesfeldt 1962, Senckenberg. Biol., Frankfurt a. Main 43(3):181. Holotype: SMF Type locality: Indefatigable Südküste, westliche Akademiebucht Indefatigable (Santa Cruz), Galápagos-Inseln. Distribution: Indefatigable (Santa Cruz) Island, Galápagos Archipelago. Amblyrhynchus cristatus mertensi Eibl-Eibesfeldt [San Cristóbal Marine Iguana] Original name: Amblyrhynchus cristatus mertensi Eibl-Eibesfeldt 1962, Senckenberg. Biol., Frankfurt a. Main 43(3):185. Holotype: SMF Type locality: etwa 3 km südwestlich der Wrack-Bucht der Insel Chatham (S. Cristobal) Chatman (Chatham [S. Cristobal]), Galápagos-Inseln. Distribution: Chatham (San Cristobal) and James (Santiago), Islands, Galápagos, Archipelago. Amblyrhynchus cristatus nanus Garman [Genovesa Marine Iguana] Original name: Amblyrhynchus nanus Garman 1892, Bull. Essex. Inst. 24:8 Holotype: BMNH = BMNH (Etheridge 1982). Type locality: Tower Island [Galápagos]. Distribution: Tower (Genovesa) Island, Galápagos Archipelago. Amblyrhynchus cristatus sielmanni Eibl-Eibesfeldt [Pinta Marine Iguana] Original name: Amblyrhynchus cristatus sielmanni Eibl-Eibesfeldt 1962, Senckenberg. Biol., Frankfurt a. Main 43(3):188. Holotype: SMF Type locality: Westküste der Insel Abingdon. Distribution: Abingdon (Pinta) Island, Galápagos Archipelago. Amblyrhynchus cristatus venustissimus Eibl-Eibesfeldt [Española Marine Iguana.] Original name: Amblyrhynchus cristatus venustissimus Eibl-Eibesfeldt 1956, Senckenberg. Biol., Frankfurt a. Main 37:90. Holotype: SMF Type locality: Nordküste der Insel Hood (Española). Distribution: Hood (Española) and Gardner Islands, Galápagos Archipelago. Brachylophus Cuvier [Fiji Iguana] Original name: Brachylophus Cuvier 1829, in Guérin-Ménville, Icon. Règ. Anim., Paris 1:9. Type species (by monotypy): Iguana fasciata Brongniart Distribution: Fiji and Tonga Islands. Comment: Based on Keogh et al. (2008) B. fasciatus is the sister species to the clade including B. vitiensis and B. bulabula; furthermore unpublished moelcular studies (Fisher et al. 2009) suggest that a fourth species may exist. Brachylophus bulabula Fisher, Harlow, Edwards, and Keogh in: Keogh, Edwards, Fisher, and Harlow [Fijian Banded Iguana] Original name: Brachylophus bulabula Fisher, Harlow, Edwards, and Keogh in: Keogh, Edwards, Fisher, and Harlow 2008, Phil. Trans. Royal Soc. B. 363(1508):3419. Holotype: CAS Type locality: Navuloa Village, Ovavlau Island, Republic of Fiji ( S, E). Distribution: Larger northwestern islands of the Viti Group of Fijian islands, including at least Ovalau, Gau, Kadavu, and Vit Levu. Comment: Although the recent description of this species clarifies the taxonomy in this genus, the correct identification of the species occuring on many islands is not yet certain. This is complicated by introductions of these iguanas onto non-native islands. Brachylophus fasciatus (Brongniart) [Lau Banded Iguana] Original name: Iguana fasciata Brongniart 1800, Bull. Soc. Philom., Paris 2:90. Holotype: none designated. Type locality: none given; Tonga, according to Keogh et al. (2008). Distribution: Lau island group of Fiji Islands, including at least Lakeba, Aiwa, Oneata, and Moce. Brachylophus vitiensis Gibbons [Fijian Crested Iguana] Original name: Brachylophus vitiensis Gibbons 1981, J. Herpet. 15(3):257. Holotype: MCZ Type locality: Yaduataba island (16 50' S; ' E), Fiji. Distribution: Fiji Island Group, found on the island of Yadua Taba and (presumably) the northern islands of the Yasawa group. Cachryx Cope [Yucatán Spiny-tailed Iguanas] Original name: Cachryx Cope 1866, Proc. Acad. Nat. Sci. Philadelphia 18:124. Type species (by monotypy): Cachryx defensor Cope Distribution: Yucatán Peninsula, Mexico. Comment: The recognition of this genus (long synonymized with Ctenosaura) is the result of molecular phylogenetics studies by Catherine Stephens et al. (in prep.). Cachryx alfredschmidti (Köhler) [Campeche Spiny-tailed Iguana] Original name: Ctenosaura alfredschmidti Köhler 1995, Salamandra 31(1):5. Holotype: SMF Type locality: 70 km östl. von Escarcega auf der Straße nach Chetumal, Campeche, Mexico. Distribution: Known only from the type locality on the Yucatán peninsula, in the Mexican state of Campeche. Cachryx defensor Cope [Yucatán Spiny-tailed Iguana] Original name: Cachryx defensor Cope 1866, Proc. Acad. Nat. Sci. Philadelphia 18:124. Syntypes: USNM [3 specimens] (Bailey 1928; de Queiroz 1995). Type locality: not given; Yucatán, by implication (de Queiroz 1995). Restricted type locality: Chichén Itzá, Yucatán, Mexico (Bailey 1928); inappropriate restriction (de Queiroz 1995). Distribution: Yucatán Peninsula from Campeche to Yucatán. Conolophus Fitzinger [Galápagos Land Iguanas] Original name: Hypsilophus (Conolophus) Fitzinger 1843, Syst. Rept., Wien 1:55. Type species (by original designation) Amblyrinchus Demarlii Duméril and Bibron 1837 = Amblyrhynchus subcristatus Gray 1831). Distribution: Galápagos Islands. Comment: Tsika et al. (2008) and Gentile et al. (2009) have demonstrated that this genus includes five evolutionarily significant units, only three of which have been described. Conolophus marthae Gentile and Snell [Galápagos Pink Land Iguana] Original name: Conolophus marthae Gentile and Snell 2009, Zootaxa, 2201:1. Holotype: A free-living adult male with a Passive Intergated Transponder # Type locality: approximately four km north of the Equator on the top of Volcan Wolf, Isla Isabela Galápagos National Park, Ecuador ( N; W, datum WGS84. Distribution: Volcan Wolf, northern Isla Isabela, Galápagos Archipelago. Comment: Additional support for the recognition of this species appeared in Tsika et al. (2008) and Gentile et al. (2009). Conolophus pallidus Heller [Santa Fe Island Iguana] Original name: Conolophus pallidus Heller 1903, Proc. Wash. Acad. Sci., 5:87. Holotype: CAS-SU Type locality: Barrington [= Santa Fe] Island, Galápagos Archipelago. Distribution: Santa Fe (= Barrington) Island, Galápagos Archipelago. Conolophus subcristatus (Gray) [Galápagos Land Iguana] Original name: Amb. [lyrhynchus] subcristatus Gray 1831, Zool. Misc., London 1831:6. Type: not located. Type locality: Galápagos? . Distribution: Galápagos Archipelago, including the islands of Santiago (= James), Santa Cruz (= Indefatigable), Isabela (= Albemarle), Fernandina (= Narborough), Baltra (= South Seymour), and Rábida (= Jervis). Comment: Tsika et al. (2008) and Gentile et al. (2009) have reported molecular evidence suggesting that some populations of C. subcristatus may deserve species status. However, the type locality of C. subcristatus is vague ( Galápagos? ; Gray 1831), and the holotype has not been located. Hence, it is not yet clear to which island the name subcristatus should be applied. Resolution of this problem may require the designation of a neotype (if the type cannot be found at the BMNH) and a simultaneous restriction of the type locality. Ctenosaura Wiegmann [Mesoamerican Spiny-tailed Iguanas] Ctenosaura Wiegmann 1828, Isis von Oken, Leipzig 21:371. Type species (by subsequent designation by Fitzinger 1843): Ctenosaura cycluroides Wiegmann 1828 = Lacerta acanthura Shaw Distribution: México and Panamá. Comment: A well-resolve phylogenetic hypothesis of all taxa in this genus is not yet available. Ctenosaura acanthura (Shaw) [Eastern Spiny-tailed Iguana] Original name: Lacerta Acanthura Shaw 1802, Gen. Zool., London 3(1):216. Holotype: BMNH XXII.20.a (Bailey 1928) = BMNH RR (Etheridge 1982). Type locality: not given. Designated type localities: California (Boulenger, 1885), in error (Smith and Taylor 1950); Mexico (Bailey 1928); Tampico, Tamaulipas, Mexico (Bailey 1928), inappropriate restriction (de Queiroz 1995). Distribution: Lowlands of eastern Mexico, from Tamaulipas southward to the Isthmus of Tehuantepec in southeastern Veracruz and eastern Oaxaca. Comment: Zarza et al. (2008) suggested that this taxon should be relegated to a subspecies of C. pectinata (see Comment under that species). Ctenosaura bakeri Stejneger [Utila Spiny-tailed Iguana] Original name: Ctenosaura bakeri Stejneger 1901, Proc. U.S. Natl. Mus. 23:467. Holotype: USNM Type locality: Utilla [sic] Island, Honduras. Distribution: Isla de Utila, Islas de la Bahía, Honduras. Ctenosaura clarki Bailey [Michoacan Spiny-tailed Iguana] Original name: Ctenosaura clarki Bailey 1928, Proc. U.S. Natl. Mus. 73(12):44. Holotype: MCZ Type locality: Ovopeo, Michoacan, Mexico. Corrected type locality: Oropeo at an elevation of about 1000 feet in the lower Tepalcatepec Valley about 8 miles south of La Huacana (Duellman and Duellman 1959). Distribution: Southwestern Mexico, in the Balsas-Tepalcatepec Basin in Michoacán. Ctenosaura conspicuosa Dickerson [San Esteban Spiny-tailed Iguana] Original name: Ctenosaura conspicuosa Dickerson 1919, Bull. Am. Mus. Nat. Hist. 41(10):461. Holotype: AMNH 5027 = USNM (Bailey 1928; Cochran 1961; de Queiroz 1995). Type locality: San Esteban Island, Gulf of California, Mexico. Distribution: Isla San Esteban, Sonora, Mexico; introduced onto Isla Cholludo, Sonora, Mexico. Comment: Ctenosaura conspicuosa was elevated from its former subspecific rank within C. hemilopha by Grismer 1999). Ctenosaura flavidorsalis Köhler and Klemmer [Yellow-backed Spiny-tailed Iguana] Original name: Ctenosaura flavidorsalis Köhler and Klemmer 1994, Salamandra 30(3):197. Holotype: SMF Type locality: 1 km südl. La Paz (750 m ü. N.N.; 14 16', 87 40'; Dpto. La Paz, Honduras). Distribution: Eastern Guatemala through northern El Salvador and southern Honduras. Ctenosaura hemilopha (Cope) [Baja California Spiny-tailed Iguana] Original name: Cyclura (Ctenosaura) hemilopha Cope 1863, Proc. Acad. Nat.Sci. Philadelphia 15:105. Syntypes: USNM 5295 [4 specimens]; one recataloged as USNM (de Queiroz 1995). Type locality: Cape St. Lucas ; near Soria Ranch, Cape San Lucas, Baja California, Mexico [USNM 5295] and San Nicolás, between Cape San Lucas and La Paz, Baja California, Mexico [USNM 69489] (Cochran 1961; de Queiroz 1995). Distribution: Baja California Sur, Mexico, from the vicinity of Loreto southward through the Cape Region, and Isla Cerralvo. Comment: As many as three subspecies have been recognized previously, but were synonymized by Grismer (1999). Ctenosaura macrolopha Smith [Sonoran Spiny-tailed Iguna] Original name: Ctenosaura hemilopha macrolopha Smith 1972, Great Basin Nat. 32(2):104. Holotype: FMNH Type locality: La Posa, San Carlos Bay, 10 mi NW Guaymas, Sonora. Distribution: Northwestern Mexico, from the vicinity of Hermosillo, Sonora, southward through the northern third of Sinaloa, and extreme western Chihuahua. Comment: This species was elevated from subspecific rank within Ctenosaura hemilopha by Grismer (1999). Ctenosaura melanosterna Buckley and Axtell [Black-chested Spiny-tailed Iguna] Original name: Ctenosaura melanosterna Buckley and Axtell 1997, Copeia 1997(1):139. Holotype: KU Type locality: 2 km south of Coyoles Central, Departmento of Yoro, Honduras. Distribution: North-central Honduras in the Río Aguan Valley and Cayos Cochinos. Ctenosaura nolascensis Smith [San Pedro Nolasco Spiny-tailed Iguana] Original name: Ctenosaura hemilopha nolascensis Smith 1972, Great Basin Nat. 32(2):107. Holotype: UCM Type locality: Isla San Pedro Nolasco, Sonora. Distribution: Isla San Pedro Nolasco, Sonora, Mexico. Comment: This species was recently elevated from subspecific rank within Ctenosaura hemilopha by Grismer (1999). Ctenosaura oaxacana Köhler and Hasbun [Oaxaca Spiny-tailed Iguana] Original name: Ctenosaura oaxacana Köhler and Hasbun 2001, Senckenberg. Biol., Frankfurt 81(1/2):260. Holotype: SMF Type locality: Tehuantepec, Estado de Oaxaca, México. Distribution: Pacific versant of the Isthmus of Tehuantepec, Estado de Oaxaca, Mexico. Ctenosaura oedirhina de Queiroz [Roatán Spiny-tailed Iguana] Original name: Ctenosaura oedirhina de Queiroz 1987, Copeia 1987(4):892. Holotype: UF Type locality: approx. 4.8 km (converted from 3 miles) west of Roatán on the path to Flowers Bay, Isla de Roatán, Departamento de las Islas de la Bahía, Honduras. Distribution: Isla de Roatán and Isla de Santa Elena, Islas de la Bahía, Honduras. Ctenosaura palearis Stejneger [Paleate Spiny-tailed Iguana] Original name: Ctenosaura palearis Stejneger 1899, Proc. U.S. Natl. Mus. 21:381. Holotype: USNM Type locality: Gualan, Guatemala. Distribution: Southeastern Guatemala in the Río Motagua Valley. Ctenosaura pectinata (Wiegmann) [Pacific or Western Spiny-tailed Iguana] Original name: Cyclura pectinata Wiegmann 1834, Herpetol. Mexicana, Berlin: 42. Syntypes: ZMB (Taylor 1969; de Queiroz 1995). Type locality: Mexico (de Queiroz 1995). Restricted type locality: Colima, Colima, Mexico (Bailey 1928); inappropriate restriction (de Queiroz 1995). Distribution: Western Mexico from north of Culican in Sinaloa southward to the Isthmus of Techuantepec in southeastern Oaxaca, including Isla Isabela and Islas de las Tres Marías, Nayarit. Comment: Zarza et al. (2008) have recently demonstrated that C. pectinata comprises at least eight distinct lineages that deserve taxonomic designation. They include: North A (Sinaloa and Nayarit; for which the name brachylophus is available); North B (coastal Southern Nayarit); North C (coastal northwestern Jalisco); Colima (for which the name brevirostris may be available); Balsas (Michoacán); Guerrero (coastal Guerrero); Oaxaca (coastal southwestern Oaxaca); and South (eastern Oaxaca north to Tamaulipas, including C. acanthura). Until specific taxonomic designations are made, we tentatively recognize C. acanthura and C. pectinata as full species. Ctenosaura quinquecarinata (Gray) [Five-keeled Spiny-tailed Iguana] Original name: Cyclura quinquecarinata Gray 1842, Zool. Misc., London 1842:59. Holotype: BMNH = BMNH RR (Etheridge 1982). Type locality: Demerara? [= Georgetown, Guyana]; in error (de Queiroz 1995); South America , in error (BMNH catalogue; de Queiroz 1995). Restricted type locality: Tehuantepec, Oaxaca, Mexico (Bailey 1928), inappropriate restriction (de Queiroz 1995); restricted to the southern portion of the distribution of C. quinquecarinata in Costa Rica and Nicaragua (Hasbun and Köhler 2001). Distribution: Nicaragua to northwestern Costa Rica. Ctenosaura similis (Gray) [Central American Spiny-tailed Iguana or Black Iguana] Original name: Iguana (Ctenosaura) Similis Gray 1831, in Griffiths, Cuvier Anim. Kingd., London 9:38. Type: Mus. [of Mr.] Bell [number not given] (de Queiroz 1995); not located (Bailey 1928). Type locality: not given. Designated type locality: Tela, Honduras, Central America (Bailey 1928), inappropriate restriction (de Queiroz 1995). Distribution: From the Isthmus de Tehuantepec southward through Central America on both versants to Panamá City and Colón, Panamá. Cyclura Harlan [Rock or Ground Iguanas] Original name: Cyclura Harlan 1824, J. Acad. Nat. Sci. Philadelphia 4:250. Type species (subsequent designation by Fitzinger, 1843): Cyclura carinata Harlan Distribution: West Indies from the Bahamas through the Greater Antilles. Comment: A well-reolved phylogenetic hypothesis was published by Malone et al. (2000). Cyclura carinata Harlan [Turks and Caicos Rock Iguana] Original name: Cyclura carinata Harlan 1824, J. Acad. Nat. Sci. Philadelphia 4:250. Type: not located (Etheridge 1982). Type locality: Turk's Island. Distribution: Turks and Caicos Islands and Booby Cay off Mayaguana in the Bahamas. Comment: Cyclura carinata bartschi Cochran 1931 (from Booby Cay off Mayaguana in the Bahamas) was recently synonymized with the nominate subspecies (Bryan et al. 2007). However, molecular work in progress (Stephen, Welch, Gerber, and Pasachnik, unpublished) suggests that populations on the western cays of the Caicos are genetically different from other populations. Cyclura collei Gray [Jamaican Rock Iguana] Original name: Cyclura Collei Gray 1845, Cat. Spec. Liz. Coll. Brit. Mus., London 190. Holotype: BMNH Type locality: Jamaica. Distribution: Jamaica, currently restricted to the Hellshire Hills in the southeast. Cyclura cornuta (Bonnaterre) [Rhinoceros Iguana] Original name: Lacerta Cornuta Bonnaterre 1789, Tab. Encycl. Méth. Règ. Nat., Erpét., Paris: 40. Type: not located (Etheridge 1982). Type locality: Sainte-Domingue dans les mornes de l'hôpital, entre L'Artibonite and les Gonaves. Distribution: Hispaniola, including Isla Beata, Isla Saona, Ile de la Gonave, Ile de la Petite Gonave, Ile Grande Cayemite, and Ile de la Tortue, and Mona Island. Comment: Most authors have included C. stejnegeri and the extinct C. onchiopis as subspecies of Cyclura cornuta, but Powell (1999), Powell and Glor (2000), and Glor et al. (2000) recommended the specific status of all three. However, Malone et al. (2000) demonstrated little genetic distance between cornuta and stejnegeri and relegated them to subspecies status. Cyclura cornuta cornuta (Bonnaterre) [Rhinoceros Iguana] Original name: Cyclura cornuta cornuta (Bonnaterre). See species account. Distribution: Hispaniola. Cyclura cornuta stejnegeri Barbour and Noble [Mona Island Iguana] Original name: Cyclura stejnegeri Barbour and Noble 1916, Bull. Mus. Comp. Zool., Cambridge 60(4):163. Holotype: USNM Type locality: Mona Island. Distribution: Isla Mona, situated between Hispaniola and Puerto Rico. Cyclura cychlura (Cuvier) [Bahaman Rock Iguana] Original name: Iguana cychlura Cuvier 1829, Règ. Anim., Ed. 2
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