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Enyalioides heterolepis (BOCOURT, 1874)

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Higher TaxaHoplocercidae, Iguania, Sauria, Squamata (lizards)
Subspecies 
Common NamesE: Bocourt's Dwarf Iguana
S: Iguana enana 
SynonymEnyalus heterolepis BOCOURT 1874: 1
Enyalius heterolepis — GÜNTHER 1885: 56
Enyalioides heterolepis — BOULENGER 1885: 114
Enyalioides insulae BARBOUR 1905: 100
Enyalioides Mocquardi DESPAX 1911
Enyalioides heterolepis — PETERS & DONOSO-BARROS 1970: 114
Enyalioides heterolepis — KÖHLER 2000: 81
Enyalioides heterolepis — TORRES-CARVAJAL et al. 2011 
DistributionNW Ecuador, Colombia (incl. Valle del Cauca, Chocó), Panama (Coclé etc.)

Type locality: Veragua, Panama.  
Reproductionoviparous 
TypesLectotype: MNHN-RA 4067
Lectotype: MNHN 4067 (= MHNP) (Guibe, 1954) 
DiagnosisDiagnosis (genus): Enyalioides can be distinguished from Hoplocercus by having a laterally compressed or rounded (rather than depressed) tail that is longer than the body (i.e., tail length > SVL). Enyalioides also differs from Hoplocercus and Morunasaurus in having pointed, conical scales on dorsal surface of head and gular region; raised scales posterior to the superciliaries; keeled dorsal scales; a continuous vertebral crest formed by large, trihedral or conical scales (except E. palpebralis, in which most specimens have a nuchal gap in the vertebral crest); and one or more enlarged pretympanic scales (except E. heterolepis). Enyalioides can be further distinguished from Morunasaurus by having a parietal eye [from TORRES-CARVAJAL et al. 2011]. ETHERIDGE 1969 presents a comparison of Enyalius and Enyalioides.


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CommentSynonymy that of PETERS & OREJAS-MIRANDA 1970.

Type species: Enyalus heterolepis BOCOURT 1874 is the type species of the genus Enyalioides BOULENGER 1885.

Key: TORRES-CARVAJAL et al. 2015: 126 and Venegas et al. 2024 provide keys to all species of the Hoplocercinae, including the species of Enyalioides.

Phylogenetics: TORRES-CARVAJAL et al. 2023 produced a phylogeny of the whole (sub-) family hoplocerinae, including Enyalioides. 
EtymologyThe species appears to be named after the heterogenous scales (Greek = “lepis” = scale or bark) on the dorsum.

The genus name is derived from the Greek Enyalius = the name of a war god and of a taxon previously confused with this one + -oides = like, resembling. 
References
  • Abuys, A. 1982. De slangen van Suriname, deel III: De familie Colubridae (algemene gegevens en het genus Chironius) / The snakes of Surinam, part III: The family Colubridae (General data and the genus Chironius). Litteratura Serpentium 2 (5): 226-245 - get paper here
  • Barbour, T. in: Bangs, Outram; Brown, Wilmont W.; Thayer, John E.; Barbour, Thomas 1905. The Vertebrata of Gorgona Island, Colombia. 1 Introduction. Mammalia. 2. Physical aspect and climate; Fauna. 3 Aves. 4 Reptilia; Amphibia. Bull. Mus. Comp. Zool. Harvard 46 (5): 87-102 - get paper here
  • Bocourt,M.F. 1874. Deux notes sur quelques sauriens de l'Amérique tropicale. Ann. Sci. Nat. Zool. Paris (5) 19 (4): 1-5 [1873] - get paper here
  • Boulenger, G.A. 1885. Catalogue of the lizards in the British Museum (Natural History). Vol. 2, Second edition. London, xiii+497 pp. - get paper here
  • Boulenger, George A. 1880. Reptiles et Batraciens recueillis par M. Emile de Ville dans les Andes de l'équateur. Bulletin de la Société Zoologique de France 5: 41-48 - get paper here
  • Carrera, C. et al. 2009. Guía de Campo de los Pequeños Vertebrados del Distrito Metropolitano de Quito (DMQ). Publicación Miscelánea N° 5. Serie de Publicaciones del Museo Ecuatoriano de Ciencias Naturales (MECN) – Fondo Ambiental del MDMQ. 1-89 pp. Imprenta Nuevo Arte. Quito-Ecuador.
  • Carvajal-Cogollo, J.E.; L.E. Rojas-Murcia. & G. Cárdenas-Arévalo 2020. Reptiles del Caribe colombiano/ Reptiles of the Colombian Caribbean. Tunja: Editorial UPTC, 268 pp. - get paper here
  • Castro-Herrera, F. & Vargas-Salinas, F. 2008. Anfibios y reptiles en el departamento del Valle del Cauca, Colombia. Biota Colombiana 9 (2): 251 - 277 - get paper here
  • CASTRO-HERRERA, FERNANDO; ANYELET VALENCIA-AGUILAR, DIEGO VILLAQUIRAN-Martínez 2012. Diversidad de Anfibios y Reptiles del Parque Nacional Natural Isla Gorgona. Universidad del Valle, Santiago de Cali, Valle del Cauca, 112 pp. - get paper here
  • Cortés Gómez, Angela María; Anyelet Valencia Aguilar, Diana Marcela Torres Domínguez, Lina María García Calderón, Diego Fernando Villaquirán Martínez, Andrea del Pilar Cáceres Franco, Fernando Castro Herrera 2010. Guía de los anfibios y reptiles. Área en conservación de la microcuenca Quebrada Pericos. [Valle de Cauca, Colombia]. Corporación Autónoma Regional del Valle del Cauca, Santiago de Cali: CVC, 37 pp. - get paper here
  • Despax, R. 1911. Note préliminaire relative aux lézards rapportés de l’Équateur par M. le Dr Rivet. Bull. Mus. natl. Hist. nat. Paris 17: 9-12 - get paper here
  • Eisenberg, Tobias and Nikola Pantchev 2010. Notes on body size and natural history of Enyalioides heterolepis (Bocourt 1874) in its northernmost population in Panama. Herpetological Bulletin (111) - get paper here
  • Etheridge, R. 1969. A review of the Iguanid lizard genus Enyalius. Bulletin of the British Museum (Natural History), Zoology 18 (8): 233-260 - get paper here
  • Fuentes, Rogemif; Aschcroft, Jesse; Erick Barría, Helio Quintero-Arrieta, Alexis Baules, Abel Batista, Eduardo Zambrano, Marcos Ponce 2023. Herpetological diversity in forests of Portobelo National Park, Colón Biological Corridor, Panama. Reptiles & Amphibians 30 (1): e18434 - get paper here
  • Günther, A. C. L. G. 1885. Reptilia and Batrachia. Biologia Centrali-Américana. Taylor, & Francis, London, 326 pp. [published in parts from 1885-1902; reprint by the SSAR 1987] - get paper here
  • Köhler, G. 2000. Reptilien und Amphibien Mittelamerikas, Bd 1: Krokodile, Schildkröten, Echsen. Herpeton Verlag, Offenbach, 158 pp.
  • Lips,K. 1999. Geographic distribution. Enyalioides heterolepis. Herpetological Review 30: 52 - get paper here
  • Nicholson, Kirsten E. 1998. Geographic distribution. Enyalioides heterolepis. Herpetological Review 29 (3): 174 - get paper here
  • Páez, V.P. et al. 2002. Guía de campo de algunas especies de anfibios y reptiles de Antioquia. Universidad de Antioquia, - get paper here
  • Pinto-Erazo, M.A., Calderón-Espinosa, M.L., Medina-Rangel, G.F. & Méndez-Galeano, M.A. 2020. Herpetofauna from two municipalities of southwestern Colombia. Biota Colombiana 21(1): 41-57. DOI: 10.21068/c2020.v21n01a04 - get paper here
  • Ríos, E.E.; C.F. Hurtado P., J.T. Rengifo M., and F. Castro Herrera. 2011. LAGARTOS EN COMUNIDADES NATURALES DE DOS LOCALIDADES EN LA REGIÓN DEL CHOCÓ DE COLOMBIA. Herpetotropicos 5 (2): 85-92 - get paper here
  • Torres-Carvajal O, Pazmiño-Otamendi G, Salazar-Valenzuela D. 2019. Reptiles of Ecuador: a resource-rich portal, with a dynamic checklist and photographic guides. Amphibian & Reptile Conservation 13 (1): [General Section]: 209–229 (e178) - get paper here
  • Torres-Carvajal O, Venegas PJ, de Queiroz K 2015. Three new species of woodlizards (Hoplocercinae, Enyalioides) from northwestern South America. ZooKeys 494: 107-132. doi: 10.3897/zookeys.494.8903 - get paper here
  • Torres-Carvajal, Omar; Fernanda P. Werneck, Igor Yuri Fernandes & Kevin de Queiroz 2023. Spiny tails and Clades: A Fully Sampled phylogeny of hoplocercine lizards (Iguanidae/ Hoplocercinae) and its taxonomic and nomenclatural Implications. Bull. Phylogenet. Nom. 001 (1): 00–02 - get paper here
  • Torres-Carvajal, Omar; RICHARD ETHERIDGE & KEVIN DE QUEIROZ 2011. A systematic revision of Neotropical lizards in the clade Hoplocercinae (Squamata: Iguania). Zootaxa 2752: 1–44 - get paper here
  • Valencia-Zuleta A, Jaramillo-Martínez AF, Echeverry-Bocanegra A, Viáfara-Vega R, Hernández-Córdoba O, Cardona-Botero VE, Gutiérrez-Zúñiga J, Castro-Herrera F. 2014. Conservation status of the herpetofauna, protected areas, and current problems in Valle del Cauca, Colombia. Amphibian & Reptile Conservation 8 (2): 1–18 (e87) - get paper here
  • Venegas PJ, Torres-Carvajal O, Duran V, de Queiroz K 2013. Two sympatric new species of woodlizards (Hoplocercinae, Enyalioides) from Cordillera Azul National Park in northeastern Peru. ZooKeys 277: 69–90. doi: 10.3897/zookeys.277.3594 - get paper here
  • Venegas, Pablo J.; Luis A. García-Ayachi, Juan C. Chávez-Arribasplata, Axel Marchelie, Santiago Bullard, Eduardo Quispe, Juan D. Valencia, Jasmín Odar, Omar Torres-Carvajal 2024. Two new species of wood lizards (Hoplocercinae: Enyalioides) from Cordillera de Colán in north-eastern Peru. Journal of Vertebrate Biology 73 (23074): 1-17 - get paper here
  • Wiens, John J. and Richard E. Etheridge 2003. Phylogenetic relationships of Hoplocercid lizards: coding and combining meristic, morphometric, and polymorphic data using step matrices. Herpetologica 59 (3): 375-398 - get paper here
 
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