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Published: 2021-03-30

Discovery of an additional piece of the large gymnophthalmid puzzle: a new genus and species of stream spiny lizard (Squamata: Gymnophthalmidae: Cercosaurinae) from the western Guiana Shield in Venezuela

Museo de Historia Natural La Salle, Fundación La Salle de Ciencias Naturales, Caracas 1050, Distrito Capital, Venezuela
Doc Frog Expeditions/CRWild, 60504, Bahía Ballena, Uvita, Costa Rica
Laboratório de Sistemática de Vertebrados, Pontifícia Universidade Católica do Rio Grande do Sul (PUCRS), Av. Ipiranga 6681, Porto Alegre, RS 90619-900, Brazil Museo Nacional de Ciencias Naturales-CSIC, C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain
Museo Nacional de Ciencias Naturales-CSIC, C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain
Laboratório de Sistemática de Vertebrados, Pontifícia Universidade Católica do Rio Grande do Sul (PUCRS), Av. Ipiranga 6681, Porto Alegre, RS 90619-900, Brazil Department of Herpetology, American Museum of Natural History, 200 Central Park West, New York, NY 10024-5102, USA
Reptilia Amazonia biodiversity Neotropics Orinoco basin phylogeny reptile systematics taxonomy


Gymnophthalmids are a highly diverse group of Neotropical lizards and its species richness is still in process of discovery. The incorporation of molecular evidence and a noticeable increase in taxon and geographic sampling in systematic studies has led to the description of numerous new genera and species of gymnophthalmids (particularly in Cercosaurinae) in recent years. Herein we describe a new genus and species of cercosaurine lizard with crocodile-like morphology, from the Venezuelan Guiana Shield on the basis of molecular phylogenetic and morphological evidence. Kataphraktosaurus gen. nov. can be readily distinguished from all other genera of Cercosaurinae by a unique combination of morphological characters that includes heterogeneous dorsal scalation with enlarged and strongly keeled scales forming two paravertebral rows, ventral and subcaudal scales imbricated and strongly keeled, large and symmetrical cephalic scales, absence of postmental scale, palpebral disc translucent and divided, tail slightly compressed, all digits clawed, and only six femoral pores (three at each hindlimb) inserted in a clump of small scales. This genus is described as monotypic and only contains Kataphraktosaurus ungerhamiltoni sp. nov., which is known from one specimen and diagnosed by the same set of aforementioned characters. The secretive habits of this species and the remoteness of the locality may explain its singleton situation. Following the International Union for Conservation of Nature’s criteria, we categorized the new species as Data Deficient.



  1. Aguirre-Peñafiel, V., Torres-Carvajal, O., Nunes, P.M.S., Peck, M.R. & Maddok, S.T. (2014) A new species of Riama Gray, 1858 (Squamata: Gymnophthalmidae) from the Tropical Andes. Zootaxa, 3866 (2), 246–260.

    Altamirano-Benavides, M., Zaher, H., Lobo, L., Grazziotin, F.G., Nunes, P.M.S. & Rodrigues, M.T. (2013) A new species of lizard genus Potamites from Ecuador (Squamata, Gymnophthalmidae). Zootaxa, 3717 (3), 345–358.

    Arèvalo, E., Davis, S.K. & Sites Jr., J.W. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43 (3), 387–418.

    Barbour, T. (1924) Two noteworthy new lizards from Panama. Proceedings of the New England Zoological Club, 9, 7–10.

    Barrio-Amorós, C.L., Fuentes, O. & Rivas, G. (2004) Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra, 40 (3–4), 183–200.

    Boulenger, G.A. (1890) First report on additions to the lizard collection in the British Museum (Natural History). Proceedings of the Zoological Society of London, 1890, 77–86.

    Boulenger, G.A. (1911) III.―Descriptions of new reptiles from the Andes of South America, preserved in the British Museum. Annals and Magazine of Natural History, 7, 19–25.

    Burt, C.E. & Burt, M.D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 227–395.

    Castoe, T.A., Doan, T.M. & Parkinson, C.L. (2004) Data partitions and complex models in bayesian analysis: The phylogeny of gymnophthalmid lizards. Systematic Biology, 53 (3), 448–469.

    Castroviejo-Fisher, S., Vilà, C., Ayarzagüena, J., Blanc, M. & Ernst, R. (2011) Species diversity of Hyalinobatrachium glassfrogs (Amphibia: Centrolenidae) from the Guiana Shield, with the description of two new species. Zootaxa, 3132 (1), 1–55.

    Doan, T.M. & Castoe, T.A. (2003) Using morphological and molecular evidence to infer species boundaries within Proctoporus bolivianus Werner (Squamata: Gymnophthalmidae). Herpetologica, 59 (3), 432–449.

    Doan, T.M. & Castoe, T.A. (2005) Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus. Zoological Journal of the Linnean Society, 143 (3), 405–416.

    Donnelly, M.A., MacCulloch, R.D., Ugarte, C.A. & Kizirian, D. (2006) A new riparian gymnophthalmid (Squamata) from Guyana. Copeia, 2006 (3), 396–403.[396:ANRGSF]2.0.CO;2

    Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–1797.

    Fang, J.M., Vásquez-Restrepo, J.D. & Daza, J.M. (2020) Filling the gaps in a highly diverse Neotropical lizard lineage: a new and endemic genus of Cercosaurinae (Squamata: Gymnophthalmidae) with the description of two new species from the Northern Andes of Colombia. Systematics and Biodiversity, 18 (5), 417–433.

    Fritts, T.H. & Smith, H.M. (1969) A new teiid lizard genus from western Ecuador. Transactions of the Kansas Academy of Science, 72 (1), 54–59.

    Fritts, T.H., Almendariz, A. & Samec, S. (2002) A new species of Echinosaura (Gymnophthalmidae) from Ecuador and Colombia with comments on other members of the genus and Teuchocercus keyi. Journal of Herpetology, 36 (3), 349–355.[0349:ANSOEG]2.0.CO;2

    Fu, J. (2000) Toward the phylogeny of the family Lacertidae–Why 4708 base pairs of mtDNA sequences cannot draw the picture. Biological Journal of the Linnean Society, 71 (2), 203–217.

    Goicoechea, N., Padial, J.M., Chaparro, J.C., Castroviejo-Fisher, S. & De la Riva, I. (2012) Molecular phylogenetics, species diversity, and biogeography of the Andean lizards of the genus Proctoporus (Squamata: Gymnophthalmidae). Molecular Phylogenetics and Evolution, 65 (3), 953–964.

    Goicoechea, N., Frost, D.R., De la Riva, I., Pellegrino, K., Sites, J., Rodrigues, M.T. & Padial, J.M. (2016) Molecular systematics of teioid lizards (Teioidea/Gymnophthalmoidea: Squamata) based on the analysis of 48 loci under tree-alignment and similarity-alignment. Cladistics, 32 (6), 624–671.

    Gorzula, S. & Señaris, J.C. (1999 [“1998”]) Contribution to the herpetofauna of the Venezuelan Guayana I. A data base. Scientia Guaianae, 8, 1–269.

    Guayasamin, J.M., Arteaga, A. & Hutter, C.R. (2018) A new (singleton) rainfrog of the Pristimantis myersi Group (Amphibia: Craugastoridae) from the northern Andes of Ecuador. Zootaxa, 4527 (3), 323–334.

    ICZN (1999) International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature, London, 306 pp.

    IUCN (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 14. Prepared by the Standards and Petitions Committee. Available from: (accessed 20 October 2020)

    Köhler, J. & Padial, J.M. (2016) Description and phylogenetic position of a new (singleton) species of Oreobates Jiménez de la Espada, 1872 (Anura: Craugastoridae) from the Yungas of Cochabamba, Bolivia. Annals of Carnegie Museum, 84 (1), 23–38.

    Köhler, G., Bohme, W. & Schmitz, A. (2004) A new species of Echinosaura (Squamata: Gymnophthalmidae) from Ecuador. Journal of Herpetology, 38 (1), 52–60.

    Kok, P.J.R. (2015) A new species of the Pantepui endemic genus Riolama (Squamata: Gymnophthalmidae) from the summit of Murisipán-tepui, with the erection of a new gymnophthalmid subfamily. Zoological Journal of the Linnean Society, 174 (3), 500–518.

    Kok, P.J.R., MacCulloch, R.D., Means, D.B., Roelants, K., Van Bocxlaer, I. & Bossuyt, F. (2012) Low genetic diversity in tepui summit vertebrates. Current Biology, 22 (15), R589–R590.

    Kok, P.J.R., Bittenbinder, M.A., van den Berg, J.K., Marques-Souza, S., Nunes, P.M.S., Laking, A.E., Teixeira Jr., M., Fouquet, A., Means, D.B., MacCulloch, R.D. & Rodrigues, M.T. (2018) Integrative taxonomy of the gymnophthalmid lizard Neusticurus rudis Boulenger, 1900 identifies a new species in the eastern Pantepui region, north-eastern South America. Journal of Natural History, 52 (13–16), 1029–1066.

    Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874.

    Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34 (3), 772–773.

    Larsson, A. (2014). AliView: a fast and lightweight alignment viewer and editor for large data sets. Bioinformatics, 30 (22), 3276–3278.

    Lehr, E., Moravec, J., Lundberg, M., Köhler, G., Catenazzi, A. & Šmid, J. (2019) A new genus and species of arboreal lizard (Gymnophthalmidae: Cercosaurinae) from the eastern Andes of Peru. Salamandra, 55 (1), 1–13.

    Lehr, E., Moravec, J. & von May, R. (2020) A new cryptic genus of terrestrial lizard (Gymnophthalmidae: Cercosaurinae) from the eastern Andes of central Peru. Salamandra, 56 (1), 1–15.

    Lim, G.S., Balke, M. & Meier, R. (2012) Determining species boundaries in a world full of rarity: singletons, species delimitation methods. Systematic Biology, 61 (1), 165–169.

    Lotzkat, S., Batista, A., Vargas, J., Hertz, A. & Köhler, G. (2012) Reptilia, Squamata, Gymnophthalmidae, Potamites apodemus (Uzzell, 1966): Distribution extension and first records from Panama. Check List, 8 (2), 302–306.

    Mamani, L., Chaparro, J.C., Correa, C., Alarcón, C., Salas, C.Y. & Catenazzi, A. (2020) A new species of Andean gymnophthalmid lizard (Squamata: Gymnophthalmidae) from the Peruvian Andes, and resolution of some taxonomic problems. Diversity, 12 (9), 361.

    Marques-Souza, S., Prates, I., Fouquet, A., Camacho, A., Kok, P.J.R., Nunes, P.M.S., Dal Vechio, F., Recoder, R.S., Mejia, N., Junior, M.T., Barrio-Amorós, C., Cassimiro, J., Lima, J.D., de Sena, M.A. & Rodrigues, M.T. (2018) Reconquering the water: evolution and systematics of South and Central American aquatic lizards (Gymnophthalmidae). Zoologica Scripta, 47 (3), 255–265.

    Moravec, J., Šmíd, J., Štundl, J. & Lehr, E. (2018) Systematics of Neotropical microteiid lizards (Gymnophthalmidae, Cercosaurinae), with the description of a new genus and species from the Andean montane forests. ZooKeys, 774, 105–139.

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: 2010 Gateway Computing Environments Workshop (GCE). IEE, New Orleans, pp. 1–8.

    Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool’s guide to PCR. Version 2.0. Department of Zoology and Kewalo Marine Laboratory, University of Hawaii, Honolulu, 45 pp.

    Parra, V., Nunes, P.M.S. & Torres-Carvajal, O. (2020) Systematics of Pholidobolus lizards (Squamata, Gymnophthalmidae) from southern Ecuador, with descriptions of four new species. ZooKeys, 954, 109–156.

    Pellegrino, K.C.M., Rodrigues, M.T., Yonenaga-Yassuda, Y. & Sites Jr, J.W. (2001) A molecular perspective on the evolution of microteiid lizards (Squamata, Gymnophthalmidae), and a new classification for the family. Biological Journal of Linnean Society, 74 (3), 315–338.

    Pellegrino, K.C.M., Brunes, T.O., Souza, S.M., Laguna, M.M., Avila-Pires, T.C.S., Hoogmoed, M.S. & Rodrigues, M.T. (2018) On the distinctiveness of Amapasaurus, its relationship with Loxopholis Cope, 1869, and description of a new genus for L. guianensis and L. hoogmoedi (Gymnophthalmoidea/Ecpleopodini: Squamata). Zootaxa, 4441 (2), 332–346.

    Pyron, R.A., Burbrink, F.T. & Wiens, J.J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13 (93), 1–53.

    Recoder, R.S., Dal Vechio, F., Marques-Souza, S., Teixeira Jr., M., Silva-Da-Silva, M., Santos-Jr., A.P., Ribeiro, S., Barrio-Amorós, C. & Rodrigues, M.T. (2018) Geographic variation and taxonomy of red-tailed Gymnophthalmus (Squamata: Gymnophthalmidae) from Amazonian savannas. Zootaxa, 4497 (1), 061–081.

    Recoder, R., Prates, I., Marques-Souza, S., Camacho, A., Nunes, P.M.S., Dal Vechio, F., Ghellere, J.M., McDiarmid, R.W. & Rodrigues, M.T. (2020) Lizards from the Lost World: two new species and evolutionary relationships of the Pantepui highland Riolama (Gymnophthalmidae). Zoological Journal of the Linnean Society, 190 (1), 271–297.

    Rojas-Runjaic, F.J.M. & Señaris, J.C. (2018) Reptiles del Escudo Guayanés de Venezuela. In: Lasso, C.A. & Señaris, J.C. (Eds.), Fauna Silvestre del Escudo Guayanés (Colombia-Venezuela). Volumen VI. Serie Editorial Fauna Silvestre Neotropical. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, D.C., pp. 179–229.

    Rojas-Runjaic, F.J.M., Castroviejo-Fisher, S. & Barrio-Amorós, C.L. (2013) First record of the Amazonian tiny tree toad Amazophrynella minuta (Melin, 1941) (Anura: Bufonidae), for Venezuela. Check List, 9 (5), 1122–1123.

    Ruibal, R. (1952) Revisionary studies of some South American Teiidae. Bulletin of the Museum of Comparative Zoology, 106 (11), 477–529.

    Saint, K.M., Austin, C.C., Donnellan, S.C. & Hutchinson, M.N. (1998) C-mos, a nuclear marker useful for squamate phylogenetic analysis. Molecular Phylogenetics and Evolution, 10 (2), 259–263.

    Sánchez-Pacheco, S.J., Torres-Carvajal, O., Aguirre-Peñafiel, V., Nunes, P.M.S., Verrastro, L., Rivas, G.A., Rodrigues, M.T., Grant, T. & Murphy, R.W. (2018) Phylogeny of Riama (Squamata: Gymnophthalmidae), impact of phenotypic evidence on molecular datasets, and the origin of the Sierra Nevada de Santa Marta endemic fauna. Cladistics, 34 (3), 260–291.

    Señaris, J.C. & Rojas-Runjaic, F.J.M. (2020) Amphibians and reptiles of Venezuelan Guayana: diversity, biogeography and conservation. In: Rull, V. & Carnaval, A. (Eds.), Neotropical diversification: Patterns and Processes. Fascinating Life Sciences. Springer, Cham, pp. 571–633.

    Sukumaran, J. & Holder, M.T. (2010) DendroPy: a Python library for phylogenetic computing. Bioinformatics, 26 (12), 1569–1571.

    Sukumaran, J. & Holder, M.T. (2017) SumTrees: Phylogenetic Tree Summarization. 4.3.0. Available from: (accessed 26 March 2020)

    Teixeira Jr, M., Dal Vechio, F., Nunes, P.M.S., Neto, A.M., Lobo, L.M., Storti, L.F., Gaiga, R.A.J., Dias, P.H.F. & Rodrigues, M.T. (2013) A new species of Bachia Gray, 1845 (Squamata: Gymnophthalmidae) from the western Brazilian Amazonia. Zootaxa, 3636 (3), 401–420.

    Torres-Carvajal, O. & Lobos, S.E. (2014) A new species of Alopoglossus lizard (Squamata, Gymnophthalmidae) from the tropical Andes, with a molecular phylogeny of the genus. ZooKeys, 410, 105–120.

    Torres-Carvajal, O. & Mafla-Endara, P. (2013) Evolutionary history of Andean Pholidobolus and Macropholidus (Squamata: Gymnophthalmidae) lizards. Molecular Phylogenetics and Evolution, 68 (2), 212–217.

    Torres-Carvajal, O., Lobos, S.E. & Venegas, P.J. (2015) Phylogeny of Neotropical Cercosaura (Squamata: Gymnophthalmidae) lizards. Molecular Phylogenetics and Evolution, 93, 281–288.

    Torres-Carvajal, O., Lobos, S.E., Venegas, P.J., Chávez, G., Aguirre-Peñafiel, V., Zurita, D. & Echevarría, L.Y. (2016) Phylogeny and biogeography of the most diverse clade of South American gymnophthalmid lizards (Squamata, Gymnophthalmidae, Cercosaurinae). Molecular Phylogenetics and Evolution, 99, 63–75.

    Torres-Carvajal, O., Venegas, P.J. & Sales-Nunes, P.M. (2020) Description and phylogeny of a new species of Andean lizard (Gymnophthalmidae: Cercosaurinae) from the Huancabamba depression. South American Journal of Herpetology, 18 (1), 13–23.

    Uetz, P., Freed, P. & Hošek, J. (2020) The Reptile Database. Available from: (accessed 20 August 2020)

    Uzzell, T.M. (1965) Teiid lizards of the genus Echinosaura. Copeia, 1965 (1), 82–89.

    Uzzell, T. (1966) Teiid lizards of the genus Neusticurus (Reptila, Sauria). Bulletin of the American Museum of Natural History, 132, 277–328.

    Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multigene datasets with character set and codon information. Cladistics, 27 (2), 171–180.

    Vásquez-Restrepo, J.D., Ibáñez, R., Sánchez-Pacheco, S.J. & Daza, J.M. (2020) Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society, 189 (1), 287–314.

    Wenzel, J.W. & Carpenter, J.M. (1994) Comparing methods: adaptive traits and tests of adaptation. In: Eggleton, P. & Vane-Wright, R.I. (Eds.), Phylogenetics and Ecology. Academic Press, London, pp. 79–101.

    Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. dissertation, The University of Texas at Austin, University of Texas libraries, Austin, Texas, 115 pp.