Abstract
Leptodeira is one of the most widespread and taxonomically problematic snake taxa in the Americas. Here we describe a new species of Leptodeira from the Andes of southern Ecuador based on morphological and molecular data. The new species is geographically close and morphologically similar to L. ornata and L. larcorum, from which it can be distinguished by having smaller dorsal body blotches, a longer tail, and shorter spines on the hemipenial body. The shortest genetic distances between the new species and its congeners are 0.02 (16S), 0.05 (cytb), and 0.18 (ND4). The new species is restricted to the Jubones River Basin in southern Ecuador, an area of endemism for other reptile species. Our phylogenetic analysis based on mitochondrial and nuclear DNA sequence data also supports recognition of the names L. larcorum (restricted to Peru) for “L. septentrionalis larcorum”, and L. ornata for populations of “L. s. ornata” from central and eastern Panama, western Colombia, and western Ecuador. However, some samples of “L. s. ornata” from Panama and Costa Rica, as well as the new species described herein, are not included within or more closely related to L. ornata, which is sister to the clade (L. bakeri, L. ashmeadii).
References
Almendáriz, A. & Carr, J.L. (2012) Lista actualizada de los anfibios y reptiles registrados en los remanentes de bosque de la Cordillera de la Costa y áreas adyacentes del suroeste del Ecuador. Revista Politécnica, 30, 184–194.
Baird, S.F. (1859) Reptiles of the Boundary. In: Emory, W.H. (Ed.), United States and Mexican Boundary Survey under the Order of Lieut. Col. W. H. Emory, Major First Cavalry, and United States Commisioner. United States Government, Washington, D.C., pp. 33–35.
Barrio-Amorós, C.L. (2019) On the taxonomy of snakes in the genus Leptodeira, with an emphasis on Costa Rica species. IRCF Reptiles & Amphibians, 26, 1–15.
Bautista, A.R. & Smith, H.M. (1992) A new chromospecies of snake (Pseudoleptodeira) from Mexico. Bulletin of the Maryland Herpetological Society, 28, 83–98.
Bocourt, M.F. (1884) Note sur quelques ophidiens nouveax, provenant de l’Amerique intertropicale. Bulletin de la Socieìteì philomathique de Paris, 8, 133–142.
Boulenger, G.A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. 3. British Museum of Natural History, London, 726 + 19 pp.
Boulenger, G.A. (1897) Description of a new snake from the Andes of Colombia. Annals and Magazine of Natural History, Series 6, 20 (120), 523.
https://doi.org/10.1080/00222939709487395
Cadle, J.E. (1984) Molecular systematics of neotropical xenodontine snakes: II. Central American xenodontines. Herpetologica, 40, 21–30.
https://doi.org/10.2307/1445144
Cadle, J.E. (1991) Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northern Perú: new species and comments on relationships and distribution patterns. Proceedings of the Academy of Natural Sciences of Philadelphia, 143, 1–96.
Campbell, J.A. (1998) Amphibians and Reptiles of northern Guatemala, the Yucatán, and Belize. University of Oklahoma Press, Norman, Oklahoma, 400 pp.
Cope, E.D. (1886) III.—Reptiles. In: Ferrari-Perez, Catalogue of animals collected by the geographical and exploring commission of the Republic of Mexico. Proceedings of the United States National Museum, 9, 182–199.
https://doi.org/10.5479/si.00963801.559.125
Cope, E.D. (1893) Second addition to the knowledge of the Batrachia and Reptilia of Costa Rica. Proceedings of the American Philosophical Society, 31, 333–347.
Daza, J.M., Smith, E.N., Páez, V.P. & Parkinson, C.L. (2009) Complex evolution in the Neotropics: the origin and diversification of the widespread genus Leptodeira (Serpentes: Colubridae). Molecular Phylogenetics and Evolution, 53, 653–667.
https://doi.org/10.1016/j.ympev.2009.07.022
de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.
https://doi.org/10.1080/10635150701701083
Duellman, W.E. (1958) A monographic study of the colubrid snake genus Leptodeira. Bulletin of the American Museum of Natural History, 114, 7–152.
Duellman, W.E. (1979) The herpetofauna of the Andes: patterns of distribution, origins, differentiation, and present communities In: Duellman, W.E. (Ed.) The South American Herpetofauna: Its Origin, Evolution, and Dispersal. Monographs of the Museum of Natural History, The University of Kansas, Lawrence, pp. 371–459.
https://doi.org/10.5962/bhl.title.3207
Duméril, A.M.C. (1853) Prodonome de la classification des reptiles ophidiens. Mémoires de l’Académie des Sciences Paris, 23, 399–536.
https://doi.org/10.5962/bhl.title.60463
Guindon, S., Dufayard, J.-F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321.
https://doi.org/10.1093/sysbio/syq010
Fitzinger, L. (1843) Systema Reptilium. Fasciculus primus, Amblyglossae. Apud Braumüller et Seidel Bibliopolas, Vienna, 106 pp.
https://doi.org/10.5962/bhl.title.4694
Günther, A. (1868) Sixth account of new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History, Series 4, 1 (6), 413–429.
https://doi.org/10.1080/00222936808695725
Günther, A. (1872) Seventh account of new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History, Series 4, 9 (49), 13–37.
https://doi.org/10.1080/002229372011951771
Günther, A. (1885–1902) Biologia Centrali-Americana. Reptilia and Batrachia. R. H. Porter, London, 326 pp.
Hallowell, E. (1845) Descriptions of reptiles from South America, supposed to be new. Proceedings of the Academy of Natural Sciences of Philadelphia, 2, 241–247.
Hallowell, E. (1861) Report upon the Reptilia of the North Pacific Exploring Expedition, under command of Capt. John Rogers. Proceedings of the Academy of Natural Sciences of Philadelphia, 12, 480–510.
Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.
https://doi.org/10.1093/molbev/mst010
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Mentjies, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.
https://doi.org/10.1093/bioinformatics/bts199
Koch, C., Venegas, P.J., Roedder, D., Flecks, M. & Boehme, W. (2013) Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of northwestern Peru and additional information on Ameiva concolor Ruthven, 1924. Zootaxa, 3745 (2), 263–295.
https://doi.org/10.11646/zootaxa.3745.2.6
Koch, C., Venegas, P.J., Cruz, R.S. & Boehme, W. (2018) Annotated checklist and key to the species of amphibians and reptiles inhabiting the northern Peruvian dry forest along the Andean valley of the Marañón River and its tributaries. Zootaxa, 4385 (1), 1–101.
https://doi.org/10.11646/zootaxa.4385.1.1
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874.
https://doi.org/10.1093/molbev/msw054
Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) Partition-Finder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701.
https://doi.org/10.1093/molbev/mss020
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, 772–773.
https://doi.org/10.1093/molbev/msw260
Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol.1. 10th Edition. Salvii, Holmiae, 824 pp.
https://doi.org/10.5962/bhl.title.542
McCranie, J.R. (2011) The Snakes of Honduras. Society for the Study of Amphibians and Reptiles, Salt Lake City, 725 pp.
Ministerio del Ambiente del Ecuador. (2012) Sistema de Clasificacion de los Ecosistemas del Ecuador Continental. Subsecretaria de Patrimonio Natural, Quito, 232 pp.
Mulcahy, D.G. (2007) Molecular systematics of neotropical cat-eyed snakes: a test of the monophyly of Leptodeirini (Colubridae: Dipsadinae) with implications for character evolution and biogeography. Biological Journal of the Linnean Society, 92, 483–500.
https://doi.org/10.1111/j.1095-8312.2007.00855.x
Mulcahy, D.G., Beckstead, T.H. & Sites Jr., J.W. (2011) Molecular systematics of the Leptodeirini (Colubroidea: Dipsadidae) revisited: Species-tree analyses and multi-locus data. Copeia, 2011, 407–417.
https://doi.org/10.1643/CH-10-058
Myers, C.W. (2011) A new genus and new tribe for Enicognathus melanauchen Jan, 1863, a neglected South American snake (Colubridae: Xenodontinae), with taxonomic notes on some Dipsadinae. American Museum Novitates, 2011, 1–33.
https://doi.org/10.1206/3715.2
Ochoa, A., Pineda, L., Crespo, P. & Willems, P. (2014) Evaluation of TRMM 3B42 precipitation estimates and WRF retrospective precipitation simulation over the Pacific–Andean region of Ecuador and Peru. Hydrology and Earth System Sciences, 18, 3179–3193.
https://doi.org/10.5194/hess-18-3179-2014
Pérez Santos, C. & Moreno, A.G. (1991) Serpientes de Ecuador. Museo Regionali di Scienze Naturali Torino, Turin, 538 pp.
Pesantes, O.S. (1994) A method for preparing the hemipenis of preserved snakes. Journal of Herpetology, 28, 93–95.
https://doi.org/10.2307/1564686
Peters, J.A. (1960) The snakes of Ecuador. A checklist and key. Bulletin of the Museum of Comparative Zoology, 122, 489–541.
Peters, W.C.H. (1866) Über neue Amphibien (Amphibolurus, Lygosoma, Cyclodus, Masticophis, Crotaphopeltis) und Fische (Diagramma, Hapalogenys) des Kgl. Zoologischen Museums. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1866, 86–96.
QGIS Development Team (2019) QGIS Geographic Information System. Open source geospatial foundation project. Available from: http://qgis.osgeo.org/ (accessed 15 October 2019)
Quintana, C., Pennington, R.T., Ulloa, C.U. & Balslev, H. (2017) Biogeographic barriers in the Andes: Is the Amotape-Huancabamba zone a dispersal barrier for dry forest plants? Annals of the Missouri Botanical Garden, 102, 542–550.
https://doi.org/10.3417/D-17-00003A
Reyes-Puig, C., Almendáriz, A. & Torres-Carvajal, O. (2017) Diversity, threat, and conservation of reptiles from continental Ecuador. Amphibian and Reptile Conservation, 11, 51–58 (e147).
Ruthven, A.G. (1936) Leptodeira bakeri, new species. Occasional Papers of the Museum of Zoology, University of Michigan, 1936, 1–2.
Sabaj, M.H. (2020) Codes for Natural History Collections in Ichthyology and Herpetology. Copeia, 108, 593–669.
https://doi.org/10.1643/ASIHCODONS2020
Salazar-Valenzuela, D., Kuch, U., Torres-Carvajal, O., Valencia, J.H. & Gibbs, H.L. (2019) Divergence of tropical pitvipers promoted by independent colonization events of dry montane Andean habitats. Journal of Biogeography, 46, 1826–1840.
https://doi.org/10.1111/jbi.13661
Schmidt, K.P. & Walker, W.F. (1943) Snakes of the Peruvian coastal region. Zoological Series of Field Museum of Natural History, 24, 297–324.
https://doi.org/10.5962/bhl.title.3848
Shreve, B. (1947) On Venezuelan reptiles and amphibians collected by Dr. H. G. Kugler. Bulletin of the Museum of Comparative Zoology, 99, 519–537.
Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57, 758–771.
https://doi.org/10.1080/10635150802429642
Torres-Carvajal, O. (2007) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76–178.
https://doi.org/10.1655/06-001.1
Torres-Carvajal, O., Pazmiño-Otamendi, G. & Salazar-Valenzuela, D. (2019) Reptiles of Ecuador: a resource-rich online portal, with dynamic checklists and photographic guides. Amphibian and Reptile Conservation, 13 (e178), 209–229.
Torres-Carvajal, O., Pazmiño-Otamendi, G. & Salazar-Valenzuela, D. (2020) Reptiles del Ecuador. Version 2020.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Available from: https://bioweb.bio/faunaweb/reptiliaweb/ (Accessed 15 June 2020).
Venegas, P.J. (2005) Herpetofauna del bosque seco ecuatorial de Peru: taxonomía, ecología y biogeografía. Zonas Áridas, 9, 9–26.
Vuilleumier, F. (1969) Pleistocene speciation in birds living in the high Andes. Nature, 223, 1179–1180.
https://doi.org/10.1038/2231179a0
Weigend, M. (2004) Additional observations on the biogeography of the Amotape-Huancabamba zone in Northern Peru: defining the south-eastern limits. Revista Peruana de Biología, 11, 127–134.