Integrative review of cis-Andean monadal coralsnakes of the genus Micrurus (Elapidae: Serpentes) reveals considerable overestimation of species diversity
Abstract
Coralsnakes of the genus Micrurus Wagler, 1824 contain approximately 89 species distributed from the United States to Argentina. The genus Micrurus comprises numerous species and presents important taxonomic challenges, particularly in lineages where morphological similarity among species complicates species delimitation. Many species have been described based on subtle morphological differences and/or inadequate molecular analyses. In this study, we focused on cis-Andean monadal coralsnakes using an integrative taxonomic approach that combines a multilocus dataset of mitochondrial (mtDNA) and nuclear (nDNA) gene fragments with morphological data, ecological niche models (ENMs), and extent of occurrence (EOO) analyses to assess species boundaries and infer phylogenetic relationships. Maximum Likelihood and Bayesian phylogenies revealed discrepancies between morphology-based taxonomy and molecular data. Among our findings, we support the junior synonymization of M. petersi Roze, 1967, M. peruvianus Schmidt, 1936, M. steindachneri (Werner, 1901), and M. tikuna Feitosa, Da Silva Jr., Pires, Zaher & Prudente, 2015, with M. ornatissimus (Jan, 1858), and of M. albicinctus Amaral, 1926, M. annellatus annellatus (Peters, 1871), M. a. balzani (Boulenger, 1898), and M. bolivianus Roze, 1967, with M. corallinus (Merrem, 1820). In contrast, our results confirm the distinctiveness of M. langsdorffi (Wagler, 1824) and M. medemi Roze, 1967 as separate evolutionary lineages, warranting their recognition as valid species. Multivariate analyses of morphometric traits showed limited differentiation among the three focal species in both sexes. Discriminant analysis of localities in environmental niche space showed an overall classification accuracy of 92.2%. The distributions of M. langsdorffi and M. ornatissimus broadly overlap, indicating sympatry, whereas both species are allopatric with respect to M. corallinus. Both occur between the Amazon and Negro rivers, whereas M. corallinus is distributed south of the Amazon River, with an isolated population in the Brazilian Atlantic Forest. These disjunctions are consistent with Wallace’s Riverine Barrier Hypothesis, suggesting river systems as major isolating factors. Micrurus medemi occurs along the eastern Andean foothills of Colombia. These results indicate that past morphology-based and/or organellar DNA-based taxonomic studies have overestimated species diversity. Our study highlights the importance of integrative taxonomy for accurately delimiting species and improving our understanding of the evolutionary history of monadal coralsnakes.
References
Aiello-Lammens M.E., Boria R.A., Radosavljevic A., Vilela B. & Anderson R.P. 2015. spThin: an R package for spatial thinning of species occurrence records for use in ecological niche models. Ecography 38 (5): 541–545. https://doi.org/10.1111/ecog.01132
Aird S.D., da Silva Jr N.J., Qiu L., Villar-Briones A., Saddi V.A., Pires de Campos M., Grau M.L. & Mikheyev A.S. 2017. Coralsnake venomics: analyses of venom gland transcriptomes and proteomes of six Brazilian taxa. Toxins 9 (6): 1–64. https://doi.org/10.3390/toxins9060187
Alkishe A., Cobos M.E., Peterson A.T. & Samy A.M. 2020. Recognizing sources of uncertainty in disease vector ecological niche models: An example with the tick Rhipicephalus sanguineus sensu lato. Perspectives in Ecology and Conservation, 18 (2): 91–102. https://doi.org/10.1016/j.pecon.2020.03.002
Amaral A. 1925. South American snakes in the collection of the United States National Museum. Proceedings of the United States National Museum 67: 1–30.
Amaral A. 1926. Ophidios de Mato Grosso. (Contribuição II para o conhecimento dos ophidios do Brasil). Commissão de Linhas Telegraphicas Estrategicas de Matto Grosso ao Amazonas. História Natural, Zoologia 84: 1–29.
Amaral A. 1930. Lista remissiva dos ophidios da regiao neotropica. Memorias do Instituto Butantan 4: 129–271.
Amaral A. 1935. Estudos sobre ophidos neotropicos. XXXIII. Novas especies de ophidios da Colombia. Memorias do Instituto Butantan 4: 219–223.
Anderson R.P., Lew D. & Peterson A.T. 2003. Evaluating predictive models of species’ distributions: criteria for selecting optimal models. Ecological Modelling 162: 211–232. https://doi.org/10.1016/S0304-3800(02)00349-6
Arevalo 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. https://doi.org/10.1093/sysbio/43.3.387
Arteaga A., Bustamante L. & Vieira J. 2024. Reptiles of Ecuador. Khamai Foundation & Tropical Herping. Quito, Ecuador.
Avila-Pires T.C. 1995. Lizards of brazilian Amazonia (Reptilia: Squamata). Zoologische Verhandelingen 299: 1–706.
Ayres J.M. & Clutton-Brock T.H. 1992. River boundaries and species range size in Amazonian primates. American Naturalist 140: 531–537.
Barve N., Barve V., Jiménez-Valverde A., Lira-Noriega A., Maher S.P., Peterson A.T., Soberón J. & Villalobos F. 2011. The crucial role of the accessible area in ecological niche modeling and species distribution modeling. Ecological Modelling 222 (11): 1810–1819. https://doi.org/10.1016/j.ecolmodel.2011.02.011
Bernarde P.S., Batista L.C., Diesel A. & Franco L. F. 2018. A remarkable new species of coralsnake of the Micrurus hemprichii species group from the Brazilian Amazon. Salamandra 54 (4): 249–258.
Bernt M., Donath A., Jühling F., Externbrink F., Florentz C., Fritzsch G., Pütz J., Middendorf M. & Stadler P. F. 2013. MITOS: improved de novo metazoan mitochondrial genome annotation. Molecular Phylogenetics and Evolution 69 (2): 313–319. https://doi.org/10.1016/j.ympev.2012.08.023
Bickford D., Lohman D.J., Sodhi N.S., Ng P.K., Meier R., Winker K., Ingram K.K. & Das I. 2007. Cryptic species as a window on diversity and conservation. Trends in Ecology and Evolution 22 (3): 148–155. https://doi.org/10.1016/j.tree.2006.11.004
Booth T.H. 2021. A problem with variable selection in a comparison of correlative and process-based species distribution models: Comments on Higgins et al., 2020. Ecology and Evolution 11 (19): 13609–13612. https://doi.org/10.1002/ece3.7496
Boulenger G.A. 1898. A list of the reptiles and batrachians collected by the late Prof. L. Balzan in Bolivia. Annali del Museo Civico di Storia Naturale di Genova (2) 19: 128–133
Boulenger G.A. 1896. Catalogue of the Snakes in the British Museum. Trustees of the British Museum (Natural History), London.
Boulenger G.A. 1902. Descriptions of new batrachians and reptiles from the Andes of Peru and Bolivia. The Annals and Magazine of Natural History (7) 10 (59): 394–402
Brauner N. & Shacham M. 1998. Role of range and precision of the independent variable in regression of data. AIChE Journal 44 (3): 603–611. https://doi.org/10.1002/aic.690440311
Caicedo-Portilla J.R. & Lynch J.D. 2015. Coral de Villavincencio. Micrurus medemi Roze, 1967. In: Morales-Betancourt M.A., Lasso C.A., Páez V.P. & Bock B.C. (eds) Libro Rojo de Reptiles de Colombia: 98–100. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Universidad de Antioquia, Bogotá.
Campbell J.A. & Lamar W.W. 2004. The Venomous Reptiles of the Western Hemisphere. Vol. 1. Comstock Publishing Associates. Ithaca, New York.
Carpenter C.C. 1953. A study of hibernacula and hibernating associations of snakes and amphibians in Michigan. Ecology 34 (1): 74–80.
Carstens B.C., Pelletier T.A., Reid N.M. & Satler J.D. 2013. How to fail at species delimitation. Molecular Ecology 22 (17): 4369–4383. https://doi.org/10.1111/mec.12413
Castoe T.A., Streicher J.W., Meik J.M., Ingrasci M.J., Poole A.W., de Koning A.P.J., Campbell J.A., Parkinson C.L., Smith E.N. & Pollock D.D. 2012. Thousands of microsatellite loci from the venomous coralsnake (Micrurus fulvius) and variability of select loci across populations and related species. Molecular Ecology Resources 12 (6): 1105–13. https://doi.org/10.1111/1755-0998.12000
Cobos M.E., Jiménez L., Nuñez-Penichet C., Romero-Alvarez D. & Simões M. 2018. Sample data and training modules for cleaning biodiversity information. Biodiversity Informatics 13: 49–50. https://doi.org/10.17161/bi.v13i0.7600
Cobos M.E., Peterson A.T., Osorio-Olvera L. & Jiménez-García D. 2019a. An exhaustive analysis of heuristic methods for variable selection in ecological niche modeling and species distribution modeling. Ecological Informatics 53: 100983. https://doi.org/10.1016/j.ecoinf.2019.100983
Cobos M.E., Peterson A.T., Barve N. & Osorio-Olvera L. 2019b. kuenm: an R package for detailed development of ecological niche models using Maxent. PeerJ 7:e6281. https://doi.org/10.7717/peerj.6281
Cope E.D. 1868. An examination of the Reptilia and Batrachia obtained by the Orton expedition to Equador and the Upper Amazon, with notes on other species. Proceedings of the Academy of Natural Sciences of Philadelphia 20: 94–140.
Criddle S. 1937. Snakes from an ant hill. Copeia 1937 (2): 142–142.
Cundall D. & Greene H.W. 2000 Feeding in snakes. In: Schwenk K. (ed.) Feeding: Form, Function and Evolution in Tetrapod Vertebrates: 293–333. Academic Press, London.
Cunha O.R.D. & Nascimento F.P.D. 1982. Ofídios da Amazônia: XVII-Revalidação de Micrurus ornatissimus (Jan, 1958) diferenciada de M. langsdorffi (Wagler, 1824) e distribuição geográfica das duas espécies (Ophidia: Elapidae). Boletim do Museu Paraense Emílio Goeldi. Nova Série Zoologia 116: 1–17.
Cunha O.R.D. & Nascimento F.P.D. 1991. Ofídios da Amazônia. XXII: Revalidação e redescrição de Micrurus albicinctus Amaral, de Rondônia, e sobre a validade de Micrurus waehrerorum Meise, do Amazonas (Ophidia: Elapidae). Boletim do Museu Paraense Emílio Goeldi. Nova Série Zoologia 7 (1): 43–52.
Da Silva Jr N.J. & Sites Jr J.W. 1999. Revision of the Micrurus frontalis complex (Serpentes: Elapidae). Herpetological Monographs: 142–194.
Da Silva Jr. N.J., Buononato M.A. & Feitosa D. 2016. As cobras-corais do Novo Mundo. Biologia, taxonomia, venenos e envenenamentos. In: Da Silva Jr N.J. (ed.) As cobras-corais do Brasil: 47–78. Editora da Pontifícia Universidade Católica da Goiás. Goiânia, Brazil.
Da Silva Jr. N.J., Feitosa D.T., Pires G.M. & Prudente A.L.C. 2021. Coralsnake diversity in Brazil. Chapter 5. In: Da Silva Jr. N.J, Porras L., Aird S. & Prudente A.L.C (eds) Advances in Coralsnake Biology with an Emphasis on South America: 201–251. Eagle Mountain Publishing. Utah, USA.
Dayrat B. 2005. Towards integrative taxonomy. Biological Journal of the Linnean Society 85 (3): 407–417. https://doi.org/10.1111/j.1095-8312.2005.00503.x
de Queiroz K.S. 2007. Species concepts and species delimitation. Systematic Biology 56 (6): 879–886. https://doi.org/10.1080/10635150701701083
DeSalle R., Egan M.G. & Siddall M. 2005. The unholy trinity: taxonomy, species delimitation and DNA barcoding. Philosophical Transactions of the Royal Society B: Biological sciences 360 (1462): 1905–1916. https://doi.org/10.1098%2Frstb.2005.1722
Di Bernardo M., Borges-Martins M. & Silva Jr N.J. 2007. A new species of coralsnake (Micrurus: Elapidae) from southern Brazil. Zootaxa 1447 (1): 1–26. https://doi.org/10.11646/zootaxa.1447.1.1
Dixon J.R., Thomas R.A. & Greene H.W. 1976. Status of the neotropical snake Rhabdosoma poeppigi Jan, with notes on variation in Atractus elaps (Günther). Herpetologica 32 (2): 221–227.
Dowling H.G. & Savage J.M. 1960. A guide to the snake hemipenis: a survey of basic structure and systematic characters. Zoologica 45: 17–28. https://doi.org/10.5962/p.203350
Duellman W.E. 1978. The biology of an equatorial herpetofauna in Amazonian Ecuador. Miscellaneous Publication. Museum of Natural History, University of Kansas 65: 1–352.
Duleba S. & Ferreira V.L. 2014. Herpetofauna associated with termite mounds in a pasture, Mato Grosso do Sul State, Brazil. Herpetological Bulletin 127: 10–16.
Duméril A.M.C., Bibron G. & Duméril A.H.A. 1854. Erpetologie generale ou, Histoire naturelle complète des reptiles. Librairie Encyclopedique de Roret, vol. 7. Paris.
Dunn E.R. 1923. Some snakes from North Western Peru. Proceedings of the Biological Society of Washington 36: 185–188.
Dunn E.R. 1954. The Coral Snake “Mimic” Problem in Panamá. Evolution 8: 97–102.
Escobar L.E., Lira-Noriega A., Medina-Vogel G. & Peterson A. 2014. Potential for spread of the white-nose fungus (Pseudogymnoascus destructans) in the Americas: Use of Maxent and NicheA to assure strict model transference. Geospatial Health 9 (1): 221–229. https://doi.org/10.4081/gh.2014.19
Feitosa D.T. 2006. Morfologia hemipeniana de 11 espécies do gênero Micrurus Wagler, 1824 na Amazônia Brasileira, com redescrição de Micrurus filiformis (Günther, 1859) e Micrurus paraensis Cunha & Nascimento, 1973 (Serpentes, Elapidae). MSc thesis. Universidad Federal do Pará. Brazil.
Feitosa D.T., Prudente A.L.C. & Lima A.C. 2007. Redescription and variation of Micrurus paraensis Cunha & Nascimento 1973 (Serpentes: Elapidae). Zootaxa 1470 (1): 35–46. https://doi.org/10.11646/zootaxa.1470.1.2
Feitosa D.T., Da Silva Jr. N.J., Pires M.G., Zaher H. & Pudente A.L. 2015. A new species of the monadal coral snake of the genus Micrurus (Serpentes, Elapidae) from western Amazon. Zootaxa 3974 (4): 538–554. https://doi.org/10.11646/zootaxa.3974.4.5
Fernandes A.M., Gonzalez J., Wink M. & Aleixo A. 2013. Multilocus phylogeography of the Wedge-billed Woodcreeper Glyphorynchus spirurus (Aves, Furnariidae) in lowland Amazonia: Widespread cryptic diversity and paraphyly reveal a complex diversification pattern. Molecular Phylogenetics and Evolution 66 (1): 270–282. https://doi.org/10.1016/j.ympev.2012.09.033
Fernandes A.M., Cohn-Haft M., Hrbek T. & Farias I.P. 2014. Rivers acting as barriers for bird dispersal in the Amazon. Revista Brasileira de Ornitologia 22 (4): 363–373. https://doi.org/10.1007/BF03544273
Fick S.E. & Hijmans R.J. 2017. WorldClim 2: new 1 km spatial resolution climate surfaces for global land areas. International Journal of Climatology 37 (12): 4302–4315. https://doi.org/10.1002/joc.5086
Flemming P.A. & Loveridge J.P. 2003. Miombo woodland termite mounds: resource islands for small vertebrates? Journal of the Zoological Society of London 259: 161–168. https://doi.org/10.1017/S0952836902003084
Frateles L.E., Tavares G., Nakamura G., da Silva Jr. N., Terribile L. & Diniz-Filho J. 2025. The interaction between the Linnean and Darwinian shortfalls affects our understanding of the evolutionary dynamics driving diversity patterns of New World coralsnakes. Journal of Biogeography 52(1): 42–54. https://doi.org/10.1111/jbi.15014
Gandolfi S.M. & Rocha C.F. 1998. Orientation of thermoregulating Tropidurus torquatus (Sauria: Tropiduridae) on termite mounds in an open area of south-eastern Brazil. Amphibia-Reptilia 19: 319–323.
Gascon C., Malcolm J.R., Patton J.L., da Silva M.N., Bogart J.P., Lougheed S.C., Peres C.A., Neckel S. & Boag P.T. 2000. Riverine barriers and the geographic distribution of Amazonian species. Proceedings of the National Academy of Sciences of the United States of America 97: 1367213677. https://doi.org/10.1073/pnas.230136397
Grismer L.L., Wood Jr P.L., Le M.D., Quah E.S. & Grismer J.L. 2020. Evolution of habitat preference in 243 species of Bent-toed geckos (Genus Cyrtodactylus Gray, 1827) with a discussion of karst habitat conservation. Ecology and Evolution 10 (24): 13717–13730. https://doi.org/10.1002/ece3.6961
Guayasamin J.M., Ribas C.C., Carnaval A.C., Carrillo J.D., Hoorn C., Lohmann L.G., Riff D., Ulloa C. & Albert J.S. 2024. Evolution of Amazonian biodiversity: A review. Acta Amazonica 54: e54bc21360.
Guisan A., Edwards Jr T.C. & Hastie T. 2002. Generalized linear and generalized additive models in studies of species distributions: setting the scene. Ecological Modelling 157 (2–3): 89–100. https://doi.org/10.1016/S0304-3800(02)00204-1
Günther A. 1868. Sixth account of new species of snakes in the collection of the British Museum. Annals and Magazine Natural History 1: 413–429. https://doi.org/10.1080/00222936808695725
Gutberlet Jr R.L. & Harvey M.B. 2004. The evolution of New World venomous snakes. In: Campbell W.L. & Lamar W.W. (eds) The Venomous Reptiles of the Western Hemisphere, Vol. 2: 634–682. Comstock Pub. Associates. Ithaca, New York.
Harvey M.B., Aparicio J. & Gonzalez L. 2003. Revision of the venomous snakes of Bolivia: Part 1. The coralsnakes (Elapidae: Micrurus). Annals of the Carnegie Museum 72: 1–52. https://doi.org/10.5962/p.215087
Hayes F. E. & Sewlal J.A. 2004. The Amazon River as a dispersal barrier to passerine birds: effects of river width, habitat and taxonomy. Journal of Biogeography 31 (11): 1809–1818.
Hoang D.T., Chernomor O., Von Haeseler A., Minh B.Q. & Vinh L.S. 2018. UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35 (2): 518–522. https://doi.org/10.1093/molbev/msx281
Hoge A.R. & Romano S.A. 1969. Espécies registradas para o Brasil (Serpentes). Ciência e Cultura 21(2): 454.
Hortal J., de Bello F., Diniz-Filho J.A., Lewinsohn T.M., Lobo J.M. & Ladle R.J. 2015. Seven shortfalls that beset large-scale knowledge of biodiversity. Annual Review of Ecology, Evolution, and Systematics 46: 523–549. https://doi.org/10.1146/annurev-ecolsys-112414-054400
Hurtado J.P., Ramirez M.V., Gómez F.J.R., Fouquet A. & Fritz U. 2021. Multilocus phylogeny clarifies relationships and diversity within the Micrurus lemniscatus complex (Serpentes: Elapidae). Salamandra 57 (2): 229–239.
IUCN. 2024. The IUCN Red List of Threatened Species. Ver. 2024-2. Available from https://www.iucnredlist.org [accessed 6 Dec. 2024].
Jan G. 1858. Plan d’une iconographie descriptive des ophidiens et description sommaire de nouvelles espèces de serpents. Revue et Magasin de Zoologie Pure et Appliquée 10: 514–527.
Jones J.C. & Oldroyd B.P. 2006. Nest thermoregulation in social insects. Advances in Insect Physiology 33: 153–191. https://doi.org/10.1016/S0065-2806(06)33003-2
Jörger K.M. & Schrödl M. 2013. How to describe a cryptic species? Practical challenges of molecular taxonomy. Frontiers in Zoology 10 (1): 1–27. https://doi.org/10.1186/1742-9994-10-59
Jowers M.J., Garcia-Mudarra J.L., Charles S.P. & Murphy J.C. 2019. Phylogeography of West Indies Coralsnakes (Micrurus): Island colonisation and banding patterns. Zoologica Scripta 48 (3): 263–276. https://doi.org/10.1111/zsc.12346
Jowers M.J., Smart U., Sánchez-Ramírez S., Murphy J.C., Gómez A., Bosque R.J., Sarker G.C., Noonan B.P., Faria J.F., Harris H.D., da Silva N.J., Prudente A.L., Weber J., Kok P.J., Rivas G.A., Jadin R.C., Sasa M., Muñoz-Mérida A., Moreno-Rueda G. & Smith E.N. 2023. Unveiling underestimated species diversity within the Central American coralsnake, a medically important complex of venomous taxa. Scientific Reports 13 (11674): 1–11. https://doi.org/10.1038/s41598-023-37734-5
Kalyaanamoorthy S., Minh B.Q., Wong T.K., Von Haeseler A. & Jermiin L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14 (6): 587–589. http://doi.org/10.1038/nmeth.4285
Katoh K. & Standley D.M. 2013. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30 (4): 772–780. https://doi.org/10.1093/molbev/mst010
Kley N.J. 2001. Prey transport mechanisms in blindsnakes and the evolution of unilateral feeding systems in snakes. American Zoologist 41: 1321–1337. https://doi.org/10.1093/icb/41.6.1321
Kley N.J. & Brainerd E.L. 1999. Feeding by mandibular raking in a snake. Nature 402: 369–370. https://doi.org/10.1038/46460
Knapp C.R. & Owens A. 2008. Nesting Behavior and the use of termitaria by the Andros Iguana (Cyclura cychlura cychlura). Journal of Herpetology 42: 46–53. https://www.jstor.org/stable/40060481
Koch C., Venegas P.J., Santa Cruz R. & Böhme 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
Kuch U. & Ayala-Varela F. 2002. First record of Micrurus peruvianus Schmidt, 1936 from Ecuador. Herpetozoa 15 (3/4): 182–183
Kumazawa Y. & Nishida M. 1993. Sequence evolution of mitochondrial tRNA genes and deep–branch animal phylogenetics. Journal of Molecular Evolution 37 (4): 380–398. https://doi.org/10.1007/bf00178868
Leigh J.W. & Bryant D. 2015. POPART: full-feature software for haplotype network construction. Methods in Ecology and Evolution 6 (9): 1110–6.
Lomonte B., Rey-Suárez P., Fernández J., Sasa M., Pla D., Vargas N., Bénard-Valle N., Sanz L., Corrêa-Netto C., Núñez V., Alape-Girón A., Alagón A., Gutiérrez J.M. & Calvete J.J. 2016. Venoms of Micrurus coral snakes: Evolutionary trends in compositional patterns emerging from proteomic analyses. Toxicon 122: 7–25. https://doi.org/10.1016/j.toxicon.2016.09.008
Lougheed S.C., Gascon C., Jones D.A., Bogart J.P., & Boag P.T. 1999. Ridges and rivers: a test of competing hypotheses of Amazonian diversification using a dart-poison frog (Epipedobates femoralis). Proceedings of the Royal Society of London. Series B: Biological Sciences 266 (1431), 1829–1835.
Maddison W.P. & Maddison D.R. 2019. Mesquite: a modular system for evolutionary analysis. Ver. 3.70. Available from https://www.mesquiteproject.org/ [accessed 12 Apr. 2026].
Marques O. & Sazima I. 2021. The natural history of New World coralsnakes. In: Silva Jr. N.J., Porras L.W., Aird S.D. & Prudente A. (eds.) Advances in Coralsnakes Biology: with an Emphasis on South America: 275–290. Eagle Mountain Publishing. Eagle Mountain, Utah.
Martins M. & Oliveira M.E. 1998. Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil. Herpetological Natural History 6 (2): 78–150.
Meise W. 1938. Eine neue Korallenschlange aus dem Amazonasgebiet. Zoologischer Anzeiger 123: 20–22.
Merrem B. 1820. Versuch eines Systems der Amphibien. Marburg. Germany.
Mizuno T. & Kojima Y. 2015. A blindsnake that decapitates its termite prey. Journal of Zoology 297 (3): 220–224. http://doi.org/10.1111/jzo.12268
Montoya-Cruz A., Mancera-García D. & Díaz-Flórez R. 2021. Defensive behavior by a Villavicencio coralsnake, Micrurus medemi Roze 1967 (Squamata: Elapidae), in Colombia. Reptiles & Amphibians 28 (3): 483–484. https://doi.org/10.17161/randa.v28i3.15719
Montoya-Cruz A., Díaz-Flórez R., & Mancera-García D. 2022. A new departmental and elevational record for the Villavicencio Coralsnake, Micrurus medemi Roze 1967 (Squamata: Elapidae), in Colombia. Reptiles & Amphibians 29 (1): 189–190. https://doi.org/10.17161/randa.v29i1.16329
Moreira L., Fenolio D., Silva H. & Silva Jr N. 2009. A preliminary list of the Herpetofauna from termite mounds of the Cerrado in the Upper Tocantins river valley. Papéis Avulsos de Zoologia 49: 183–189. https://doi.org/10.1590/S0031-10492009001500001
Nagy Z. T., Dekoninck W., De Block F., Yeo K., Silue K.S. & Delsinne T. 2017. Oviposition of the snake Thelotornis kirtlandii in a parabiotic ant nest. Salamandra 53(1): 167–170.
Nascimento L.R., Silva Jr N.J., Feitosa D.T. & Prudente A.L. 2019. Taxonomy of the Micrurus spixii species complex (Serpentes, Elapidae). Zootaxa 4668 (3): 370–392. https://doi.org/10.11646/zootaxa.4668.3.4
Nascimento L.R., Graboski R., Silva Jr. N.J. & Prudente A.L. 2024. Integrative taxonomy of Micrurus ibiboboca (Merrem, 1820) (Serpentes, Elapidae) reveals three new species of coral snake. Systematics and Biodiversity 22 (1): 2315958. https://doi.org/10.1080/14772000.2024.2315958
Niceforo M.H. 1942. Los ofidios de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 5 (17): 84–101.
Nogueira C.C., Argôlo A.J., Arzamendia V., Azevedo J.A., Barbo F.E., Bérnils R.S., Bolochio B., Borges-Martins M., Brasil-Godinho M., Braz H., Buononato M., Cisneros-Heredia D., Colli G., Costa H., Franco F., Giraudo A., Gonzalez R., Guedes T., Hoogmoed M., Marques O. & Martins M. 2019. Atlas of Brazilian snakes: verified point-locality maps to mitigate the Wallacean shortfall in a megadiverse snake fauna. South American Journal of Herpetology 14: 1–274. https://doi.org/10.2994/SAJH-D-19-00120.1
Noonan B.P. & Chippindale P.T. 2006. Vicariant origin of Malagasy reptiles supports Late Cretaceous Antarctic land bridge. The American Naturalist 168 (6): 730–741. https://doi.org/10.1086/509052
Olson D.M., Dinerstein E., Wikramanayake E.D., Burgess N.D., Powell G.V., Underwood E.C., D’Amico J.A., Itoua I., Strand H.E., Morrison J.C., Loucks C.J., Allnutt T.F., Ricketts T.H., Kura Y., Lamoreux J.F., Wettengel W.W., Hedao P. & Kassem K.R. 2001. Terrestrial ecoregions of the world: a new map of life on Earth. Bioscience 51 (11): 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
Osorio-Olvera L., Lira-Noriega A., Soberón J., Peterson A.T., Falconi M., Contreras-Díaz R.G., Martínez-Meyer E., Barve V. & Barve N. 2020. ntbox: An r package with graphical user interface for modelling and evaluating multidimensional ecological niches. Methods in Ecology and Evolution 11 (10): 1199–1206. https://doi.org/10.1111/2041-210X.13452
Padial J.M., Miralles A., De la Riva I. & Vences M. 2010. The integrative future of taxonomy. Frontiers in Zoology 7 (1): 1–14. https://doi.org/10.1186/1742-9994-7-16
Palumbi S.R. 1996. Nucleic acids II: The polymerase chain reaction. In: Hillis D.M., Moritz C. & Mable B.K. (eds.) Molecular Systematics: 205–207. Sinauer Associates, Massachusetts.
Parkinson C.L., Campbell J.A. & Chippendale P.T. 2002. Multigene phylogenetic analysis of pitvipers with comments on the biogeographical history of the group. In: Schuett G.W., Hoggren M., Douglas M.E. & Greene H.W. (eds) Biology of the Vipers: 93–110. Eagle Mountain Publishing, Utah.
Patton J.L., Da Silva M.N.F. & Malcolm J.R. 1994. Gene genealogy and differentiation among arboreal spiny rats (Rodentia: Echimyidae) of the Amazon basin: a test of the riverine barrier hypothesis. Evolution 48 (4): 1314–1323.
Passos P. & Fernandes D.S. 2005. Variation and taxonomic status of the aquatic coral snake Micrurus surinamensis (Cuvier, 1817) (Serpentes: Elapidae). Zootaxa 953 (1): 1–8. https://doi.org/10.11646/zootaxa.953.1.1
Peres C.A., Patton J.L., & da Silva M. 1996. Riverine barriers and gene flow in Amazonian saddle-back tamarins. Folia Primatologica 67 (3): 113–124.
Peters J.A. & Orejas-Miranda B. 1970. Catalogue of the Neotropical Squamata. Part I. Snakes. Bulletin of the United States National Museum 297: 1–347.
Peters W. 1871. Über eine von Hrn. Dr. Robert Abendroth in dem Hochlande von Peru gemachte Sammlung von Amphibien, welche derselbe dem Königl. zoologischen Museum geschenkt hat. Monatsberichte der Königlichen Preussischen Akademie der Wissenschaften zu Berlin: 397–404.
Peterson A.T. & Soberón J. 2012. Species distribution modeling and ecological niche modeling: getting the concepts right. Natureza & Conservação 10 (2): 102–107. http://dx.doi.org/10.4322/natcon.2012.019
Pires A.C. & Marinoni L. 2010. DNA barcoding and traditional taxonomy unified through integrative taxonomy: a view that challenges the debate questioning both methodologies. Biota Neotropica 10: 339–346. https://doi.org/10.1590/S1676-06032010000200035
Pires G.M., Da Silva Jr N.J., Feitosa T.D., Prudente A., Filho G.A. & Zaher H. 2014. A new species of triadal coral snake of the genus Micrurus Wagler, 1824 (Serpentes: Elapidae) from northeastern Brazil. Zootaxa 3811 (4): 569–584. https://doi.org/10.11646/zootaxa.3811.4.8
Pisani R.G. 2009. Use of an active ant nest as a hibernaculum by small snake species. Transactions of The Kansas Academy of Science 112: 113–118. https://doi.org/10.1660/062.112.0215
Phillips S.J., Anderson R.P. & Schapire R.E. 2006. Maximum entropy modeling of species geographic distributions. Ecological Modelling 190 (3-4): 231–259. https://doi.org/10.1016/j.ecolmodel.2005.03.026
Puillandre N., Brouillet S. & Achaz G. 2021. ASAP: assemble species by automatic partitioning. Molecular ecology resources, 21(2), 609–620. https://doi.org/10.1111/1755-0998.13281
Quilumbaquín W., Carrera-Gonzalez A. & Ortega-Andrade H.M. 2023. Environmental DNA and visual encounter surveys for amphibian biomonitoring in aquatic environments of the Ecuadorian Amazon. PeerJ 11: e15455. https://doi.org/10.7717/peerj.15455
R Core Team. 2022. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from https://www.R-project.org/
Rambaut A., Drummond A.J., Xie D., Baele G. & Suchard M.A. 2018. Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67 (5): 901–904. https://doi.org/10.1093/sysbio/syy032
Rannala B. 2015. The art and science of species delimitation. Current Zoology 61 (5): 846–853. https://doi.org/10.1093/czoolo/61.5.846
Reyes-Velasco J., Adams R.H., Boissinot S., Parkinson C.L., Campbell J.A., Castoe T.A. & Smith E.N. 2020. Genome-wide SNPs clarify lineage diversity confused by coloration in coralsnakes of the Micrurus diastema species complex (Serpentes: Elapidae). Molecular Phylogenetics and Evolution 147: 106770. https://doi.org/10.1016/j.ympev.2020.106770
Ronquist F., Teslenko M., Van Der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematics Biology 61: 539–542. http://dx.doi.org/10.1093/sysbio/sys029
Rozas J., Ferrer-Mata A., Sánchez-DelBarrio J.C., Guirao-Rico S., Librado P., Ramos-Onsins S.E., & Sánchez-Gracia A. 2017. DnaSP 6: DNA sequence polymorphism analysis of large data sets. Molecular Biology and Evolution 34 (12): 3299–3302. https://doi.org/10.1093/molbev/msx248
Roze J.A. 1966. On the synonymy and type specimens of the coral snakes Micrurus corallinus and M. ibiboboca (Marcgravii). Copeia 1966 (2): 369–371.
Roze J.A. 1967. A check list of the New World venomous coralsnakes (Elapidae), with descriptions of new forms. American Museum Novitates 2287: 1–60.
Roze J.A. 1983. New World coral snakes (Elapidae): a taxonomic and biological summary. Memorias do Instituto Butantan 46: 305–338.
Roze J.A. 1989. New species and subspecies of coralsnakes, genus Micrurus (Elapidae), with notes on type specimens of several species. American Museum Novitates 2932: 1–15.
Roze J.A. 1994. Notes on taxonomy of venomous coral snakes (Elapidae) of South America. Bulletin of the Maryland Herpetological Society 30: 177–185.
Roze J.A. 1996. Coral Snakes of the Americas: Biology, Identification, and Venoms. Krieger Publishing Company, Malabar, Florida.
Sahlin K., Lim M.C. & Prost S. 2021. NGSpeciesID: DNA barcode and amplicon consensus generation from long-read sequencing data. Ecology and Evolution 11 (3): 1392–1398. https://doi.org/10.1002/ece3.7146
Samadi S. & Barberousse A. 2006. The tree, the network, and the species. Biological Journal of the Linnean Society 89 (3): 509–521. https://doi.org/10.1111/j.1095-8312.2006.00689.x
Santos C.D., Sarmento H., de Miranda F.P., Henrique-Silva F. & Logares R. 2020. Uncovering the genomic potential of the Amazon River microbiome to degrade rainforest organic matter. Microbiome 8: 151. https://doi.org/10.1186/s40168-020-00930-w
Savage J.M. 1960. A revision of the Ecuadorian snakes of the colubrid genus Atractus. Miscellaneous Publications Museum of Zoology, University of Michigan 112: 1–86.
Savage J.M. & Slowinski J.B. 1990. Short note: A simple consistent terminology for the basic colour patterns of the venomous coral snakes and their mimics. Herpetological Journal 1: 530–532.
Savage J.M. & Slowinski J.B. 1992. The colouration of the venomous coral snakes (family Elapidae) and their mimics (families Aniliidae and Colubridae). Biological Journal of the Linnean Society 45 (3): 235–254. https://doi.org/10.1111/j.1095-8312.1992.tb00642.x
Schmidt K.P. 1936. Preliminary account of coral snakes of South America. Field Museum of Natural History Publications, Zoological Series 20: 189–203.
Schmidt K.P. 1954. The annellated coral snake, Micrurus annellatus Peters. Fieldiana, Zoology 34 (30): 319–325.
Schmidt K.P. 1955. Coral snakes of the genus Micrurus in Colombia. Fieldiana, Zoology 34: 337–359.
Schlick-Steiner B.C., Steiner F.M., Seifert B., Stauffer C., Christian E. & Crozier R.H. 2010. Integrative taxonomy: a multisource approach to exploring biodiversity. Annual Review of Entomology 55: 421–438. https://doi.org/10.1146/annurev-ento-112408-085432
Silva-Haad J.J. 1994. Las Micrurus de la Amazonia colombiana: biología y toxicología experimental de sus venenos. Colombia Amazónica 7: 41–138.
Sites Jr J.W. & Marshall J.C. 2004. Operational criteria for delimiting species. Annual Review of Ecology, Evolution, and Systematics 35: 199–227. https://doi.org/10.1146/annurev.ecolsys.35.112202.130128
Shine R. & Madsen T. 1997. Prey abundance and predator reproduction: rats and pythons on a tropical Australian floodplain. Ecology 78 (4): 1078–1086. https://doi.org/10.1890/0012-9658(1997)078[1078:PAAPRR]2.0.CO;2
Slowinski J.B. 1995. A phylogenetic analysis of the New World coralsnakes (Elapidae: Leptomicrurus, Micruroides, and Micrurus) based on allozymic and morphological characters. Journal of Herpetology 29 (3): 325–338.
Slowinski J.B. & Keogh J.S. 2000. Phylogenetic relationships of elapid snakes based on cytochrome b mtDNA sequences. Molecular Phylogenetics and Evolution 15 (1): 157–164. https://doi.org/10.2307/1564981
Soini P. 1974. Polychromatism in a population of Micrurus langsdorffi. Journal of Herpetology 8 (3): 267–269. https://doi.org/10.2307/1563184
Streicher J.W., McEntee J.P., Drzich L.C., Card D.C., Schield D.R., Smart U., Parkinson C.L., Jezkova T., Smith E.N. & Castoe T.A. 2016. Genetic surfing, not allopatric divergence, explains spatial sorting of mitochondrial haplotypes in venomous coralsnakes. Evolution 70 (7): 1435–1449. https://doi.org/10.1111/evo.12967
Syfert M.M., Smith M.J. & Coomes D.A. 2013. The effects of sampling bias and model complexity on the predictive performance of MaxEnt species distribution models. PloS ONE 8 (2): e55158. https://doi.org/10.1371/journal.pone.0055158
Tamura K., Stecher G. & Kumar S. 2021. MEGA11: Molecular Evolutionary Genetics Analysis version 11. Molecular Biology and Evolution 38 (7): 3022–3027. https://doi.org/10.1093/molbev/mst197
Terribile L.C., Diniz-Filho J.A., Rodríguez M.Á. & Rangel T.F. 2009. Richness patterns, species distributions and the principle of extreme deconstruction. Global Ecology and Biogeography 18 (2): 123–136. https://doi.org/10.1111/j.1466-8238.2008.00440.x
Terribile L.C., Feitosa D.T., Pires M.G., de Almeida P.C., de Oliveira G., Diniz-Filho J.A. & da Silva Jr N. 2018. Reducing Wallacean shortfalls for the coralsnakes of the Micrurus lemniscatus species complex: Present and future distributions under a changing climate. PloS One 13 (11): e0205164. https://doi.org/10.1371/journal.pone.0205164
Terribile L.C., de Oliveira G., Lima-Ribeiro M.S., Feitosa D.T., Giraudo A.R., Martins M.M., Arzamendia V., Pires M.G., da Silva Jr N., Moura J.C., Franco F.L., Silveira R. & Diniz-Filho J.A. 2021. Macroecology and conservation of South American coralsnakes. In: da Silva Jr N., Hoggren M., Porras W.E., Aird S.A. & Prudente L. (eds) Advances in Coralsnake Biology: with an Emphasis on South America: 313–332. Eagle Mountain Publishing, Utah.
Townsend T.M., Alegre R.E., Kelley S.T., Wiens J.J. & Reeder T.W. 2008. Rapid development of multiple nuclear loci for phylogenetic analysis using genomic resources: an example from squamate reptiles. Molecular Phylogenetics and Evolution 47 (1): 129–142.
Trifinopoulos J., Nguyen L.T., von Haeseler A. & Minh B.Q. 2016. W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nuclei Acids Research 44 (W1): W232–W235. https://doi.org/10.1093/nar/gkw256
Valencia J.H. 2007. First record of the coral snake Micrurus margaritiferus Roze, 1967 in Ecuador. Herpetozoa 19 (3/4): 186–187.
Valencia J.H., Garzón-Tello K. & Barragán-Paladines M.E. 2016. Serpientes venenosas de Ecuador: Sistemática, taxonomía, historia natural, conservación, envenenamiento y aspectos antropológicos. Fundación Herpetológicas Gustavo Orcés/ Universidad de Texas, Arlington/Fondo Ambiental Nacional. Quito.
Vaser R., Sović I., Nagarajan N. & Šikić M. 2017. Fast and accurate de novo genome assembly from long uncorrected reads. Genome Research 27 (5): 737–746. https://doi.org/10.1101/gr.214270.116
Vaz-Ferreira R., Covelo de Zolessi L. & Achaval F. 1970. Oviposición y desarrollo de ofidios y lacertilios en hormigueros de Acromyrmex. Physis 29 (79): 431–459.
Vaz-Ferreira R., Covelo de Zolessi L. & Achaval F. 1974. Oviposición y desarrollo de ofidios y lacertilios en hormigueros de Acromyrmex. II. Trabajos del V Congreso Latinoamericano de Zoología 1: 232–244.
Vences M., Miralles A. & Dufresnes C. 2024. Next-generation species delimitation and taxonomy: Implications for biogeography. Journal of Biogeography 51 (9): 1–14. https://doi.org/10.1111/jbi.14807
Venables W.N. & Ripley B.D. 2002. Modern Applied Statistics with S, 4th Edition. Springer, New York.
Wagler J. 1824. Serpentum Brasiliensium species novae = ou, Histoire naturelle des espèces nouvelles de serpens: and recueillies et observées pendant le voyage dans l’intérieur du Brésil dans les années 1817, 1818, 1819, 1820. Monachii Typis F.S. Hübschmanni, Munich.
Wallace A.R. 1852. On the monkeys of the Amazon. Annals and Magazine of Natural History 14 (84): 451–454. https://doi.org/10.1080/037454809494374
Werner F. 1901. Über Reptilien und Batrachier aus Ecuador und Neu-Guinea. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 51: 593–614.
Werner F. 1927. Neue oder wenig bekannte Schlangen aus dem Wiener naturhistorischen Staatsmuseum. (III Teil). Sitzungsberichte der Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Klasse 135: 243–257.
Wied-Neuwied M. 1820. Über die Cobra Coral oder Cobras Coraes der Brasilianer. Nova Acta Academiae Caesareae Leopoldino-Carolinae 10: 105–110.
Will K.W., Mishler B.D. & Wheeler Q.D. 2005. The perils of DNA barcoding and the need for integrative taxonomy. Systematic Biology 54 (5): 844–851. https://doi.org/10.1080/10635150500354878
Zaher H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History 240: 1–168.
Zhang J., Kapli P., Pavlidis P. & Stamatakis A. 2013. A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29 (22): 2869–2876. https://doi.org/10.1093/bioinformatics/btt499
Copyright (c) 2026 Jorge H. Valencia, Eric N. Smith, Biraj Shrestha, Juan C. Chávez-Arribasplata, Pablo J. Venegas, Walter Quilumbaquin, Alejandro Espinosa de los Monteros, Omar Torres-Carvajal, Juan C. Chaparro, Andrés Lira-Noriega, H. Mauricio Ortega-Andrade

This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT PERMITTED to post their submitted work online (e.g., in institutional repositories or on personal websites) prior to publication, as it may lead to nomenclatural issues.
