Submitted:
15 July 2026
Posted:
16 July 2026
You are already at the latest version
Abstract
Keywords:
1. Introduction
2. Endless Forms Most Beautiful: The Known Diversity of Ecuadorian Scorpions
2.1. Bothriuridae
2.2. Buthidae
2.3. Caraboctonidae
2.4. Chactidae
2.5. Troglotayosicidae
3. Towards Integrative Thinking in Ecuadorian Scorpion Biodiversity Research
4. Understanding and Fighting the Burden of Scorpionism Across Ecuador
5. The Toxinology of Ecuadorian Scorpions
6. Research Priorities for Combating Scorpionism in Ecuador
Acknowledgments
References
- Dionisio-da-Silva, W.; Foerster, S.Í.A.; Gallão, J.E.; Lira, A.F.A. What’s for dinner? Prey consumption by Neotropical scorpions across contrasting environments. J. Arachnol. 2024, 52, 26–30. [Google Scholar] [CrossRef]
- Polis, G.A. The biology of Scorpions; Stanford University Press, 1990. [Google Scholar]
- Simone, Y.; Meijden, A.V.D. Armed stem to stinger: a review of the ecological roles of scorpion weapons. J. Venom. Anim. Toxins Trop. Dis. 2021, 27, e20210002. [Google Scholar] [CrossRef] [PubMed]
- Tobassum, S.; Tahir, H.M.; Arshad, M.; Zahid, M.T.; Ali, S.; Ahsan, M.M. Nature and applications of scorpion venom: an overview. Toxin Rev. 2020, 39, 214–225. [Google Scholar] [CrossRef]
- Ghosh, A.; Roy, R.; Nandi, M.; Mukhopadhyay, A. Scorpion Venom–Toxins that Aid in Drug Development: A Review. Int. J. Pept. Res. Ther. 2019, 25, 27–37. [Google Scholar] [CrossRef] [PubMed]
- Rein, J. O. The Scorpion Files. The Scorpion Files [Internet]. 2017. Available online: https://www.ntnu.no/ub/scorpion-file.
- Borges, A. Scorpionism and Dangerous Scorpions in Central America and the Caribbean Region. In Scorpion Venoms: Scorpion Venoms; Gopalakrishnakone, P., Ed.; Springer Netherlands: Dordrecht, 2021; pp. 1–27. [Google Scholar] [CrossRef]
- Borges, A.; Graham, M.R. Phylogenetics of Scorpions of Medical Importance. In Venom Genomics and Proteomics; Gopalakrishnakone, P., Calvete, J.J., Eds.; Springer Netherlands: Dordrecht, 2016; pp. 1–20. [Google Scholar] [CrossRef]
- Lourenço, W.R. The evolution and distribution of noxious species of scorpions (Arachnida: Scorpiones). J. Venom. Anim. Toxins Trop. Dis. 2018, 24, 1. [Google Scholar] [CrossRef] [PubMed]
- Ward, M.J.; Ellsworth, S.A.; Nystrom, G.S. A global accounting of medically significant scorpions: Epidemiology, major toxins, and comparative resources in harmless counterparts. Toxicon 2018, 151, 137–155. [Google Scholar] [CrossRef] [PubMed]
- Cid-Uribe, J.I.; Veytia-Bucheli, José Ignacio; Romero-Gutierrez, Teresa; Ortiz, Ernesto; Possani, L.D. Scorpion venomics: a 2019 overview. Expert Rev. Proteom. 2020, 17, 67–83. [Google Scholar] [CrossRef] [PubMed]
- Chippaux, J.-P.; Goyffon, M. Epidemiology of scorpionism: A global appraisal. Acta Trop. 2008, 107, 71–79. [Google Scholar] [CrossRef] [PubMed]
- Isbister, G.K.; Bawaskar, H.S. Scorpion Envenomation. N Engl. J. Med. 2014, 371, 457–463. [Google Scholar] [CrossRef] [PubMed]
- Monteiro, W.M.; Gomes, J.; Fé, N.; Mendonça da Silva, I.; Lacerda, M.; Alencar, A.; et al. Perspectives and recommendations towards evidence-based health care for scorpion sting envenoming in the Brazilian Amazon: A comprehensive review. Toxicon 2019, 169, 68–80. [Google Scholar] [CrossRef] [PubMed]
- Hernández-Muñoz, E.A.; Borges, A.; Zavala-Sánchez, E.V.; de Arias, A.R.; Oukkache, N.; de Souza, C.M.V.; et al. Scorpion sting envenomation: a neglected tropical disease in the shadow of global health priorities: an urgent call to action. BMJ Glob. Health 2025, 10. [Google Scholar] [CrossRef] [PubMed]
- Gutiérrez, J.M.; Chippaux, J.P.; Isbister, G.K. PLOS Neglected Tropical Diseases broadens its coverage of envenomings caused by animal bites and stings. PLoS Negl. Trop. Dis. 2021, 15, e0009481. [Google Scholar] [CrossRef] [PubMed]
- Hernández-Muñoz, E.A.; Zavala-Sánchez, E.V. Scorpion sting envenomation: should it be considered a neglected tropical disease? Int. J. Epidemiol. 2024, 53, dyae070. [Google Scholar] [CrossRef] [PubMed]
- Borges, A.; De Sousa, L.; Borja-Cabrera, G.P.; Rivera, A. Scorpion envenoming in Ecuador: district-level risk, updated scorpion diversity, and challenges for treatment and public health. Acta Trop. 2026, 273, 107942. [Google Scholar] [CrossRef] [PubMed]
- Vasconez-Gonzalez, J.; Izquierdo-Condoy, J.S.; Miño, C.; de Lourdes Noboa-Lasso, M.; Ortiz-Prado, E. Epidemiological and geodemographic patterns of scorpionism in Ecuador: A nationwide analysis (2021–2024). Toxicon X 2025, 26, 100218. [Google Scholar] [CrossRef] [PubMed]
- Brito, G.; Borges, A. A checklist of the scorpions of Ecuador (Arachnida: Scorpiones), with notes on the distribution and medical significance of some species. J. Venom. Anim. Toxins Trop. Dis. 2015, 21, 23. [Google Scholar] [CrossRef] [PubMed]
- Kleemann, J.; Koo, H.; Hensen, I.; Mendieta-Leiva, G.; Kahnt, B.; Kurze, C.; et al. Priorities of action and research for the protection of biodiversity and ecosystem services in continental Ecuador. Biol. Conserv. 2022, 265, 109404. [Google Scholar] [CrossRef]
- Lourenço, W.R. Diversity and endemism in tropical versus temperate scorpion communities. Biogeographica 1994, 70, 155–160. [Google Scholar]
- Arbuckle, K. Evolutionary Context of Venom in Animals. In Evolution of Venomous Animals and Their Toxins; Springer: Dordrecht, 2017; pp. 3–31. [Google Scholar] [CrossRef]
- Bordon K de, C.F.; Cologna, C.T.; Fornari-Baldo, E.C.; Pinheiro-Júnior, E.L.; Cerni, F.A.; Amorim, F.G.; et al. From Animal Poisons and Venoms to Medicines: Achievements, Challenges and Perspectives in Drug Discovery. Front Pharmacol. 2020, 11. [Google Scholar] [CrossRef] [PubMed]
- Surm, J.M.; Moran, Y. Insights into how development and life-history dynamics shape the evolution of venom. EvoDevo 2021, 12, 1. [Google Scholar] [CrossRef] [PubMed]
- Cekalovic, K. T. Brachistosternus pegnai n. sp. de escorpión para el Ecuador (Scorpionida-Bothriuridae). In Bol Soc Biológica Concepc; 1969; pp. 163–171. [Google Scholar]
- Cekalovic, K. T.; Artigas, J.N. Catálogo de los tipos depositados en la colección del Departamento de Zoología de la Universidad de Concepción, Chile : parte III. (Hasta Septiembre de 1976). In Bol Soc Biológica Concepc.; 1981; pp. 75–107. [Google Scholar]
- Ojanguren-Affilastro, A.A.; Mattoni, C.I.; Ochoa, J.A.; Ramírez, M.J.; Ceccarelli, F.S.; Prendini, L. Phylogeny, species delimitation and convergence in the South American bothriurid scorpion genus Brachistosternus Pocock 1893: Integrating morphology, nuclear and mitochondrial DNA. Mol. Phylogenet Evol. 2016, 94, 159–170. [Google Scholar] [CrossRef] [PubMed]
- Lourenço, W.R. Scorpions cavernicoles de l’Équateur: Tityus demangei n. sp. et Ananteris ashmolei n. sp. (Buthidae); Troglotayosicus vachoni n. gen., n. sp. (Chactidae), scorpion troglobie. In Bull Muséum Natl D’Histoire Nat Sect Zool Biol Écologie Anim; 1981; pp. 635–662. [Google Scholar]
- Lourenço, W.R. Révision du genre Ananteris Thorell, 1891 (Scorpiones, Buthidae) et description de six espèces nouvelles. Bull. Muséum Natl. Hist. Nat. 1982, 4, 119--151. [Google Scholar] [CrossRef]
- Lourenço, W.R. Some remarks about Ananteris festae Borelli, 1899 and description of a new species of Ananteris Thorell from Ecuador (Scorpiones, Buthidae). Entomol. Mitt. Zool. Mus. Hambg. 1999, 13, 95–100. [Google Scholar]
- Ythier, E. A new high-altitude scorpion species of the genus Ananteris Thorell, 1891 (Scorpiones: Ananteridae) from the Pico da Neblina, Brazil. 2024, 12, 1–9. [Google Scholar] [CrossRef] [PubMed]
- Prendini, L. All genera of the world: Order Scorpiones (Animalia: Arthropoda: Arachnida). Megataxa 2026, 19, 270–378. [Google Scholar] [CrossRef]
- Behr, H.; Correa, O. Identificacion de los alacranes de la familia Buthidae en la Urbanización Puerto Azul. Guayaquil. Rev. Ecuat. Hig. Med. Trop. 198737, 71–76. [Google Scholar]
- Campos, R.F. Breve contribucion al conocimiento de los Escorpionidos (Alacranes) del Ecuador. Rev. Col. Nac. Vicente Rocafuerte 193113, 117–125. [Google Scholar]
- de Armas, L.F.; Teruel, R.; Kovařík, F. On Centruroides margaritatus (Gervais, 1841) and closely related species (Scorpiones: Buthidae). Euscorpius 2011, 1–16. [Google Scholar] [CrossRef]
- Lourenço, W.R.; Ythier, E. The remarkable scorpion diversity in the Ecuadorian Andes and description of a new species of Tityus C. L. Koch, 1836 (Scorpiones, Buthidae); ZooKeys, 2013; pp. 1–13. [Google Scholar] [CrossRef] [PubMed]
- Ythier, E. Two new species of Hadruroides Pocock, 1893 from Peru and Ecuador (Scorpiones, Caraboctonidae). Faunitaxys 2021, 9, 1–8. [Google Scholar]
- Lourenço, W.R. Les scorpions (Chelicerata, Scorpiones) de l’Equateur, avec quelques considérations sur la biogéographie et la diversité des espèces. Rev. Suisse Zool. 1995, 102, 61–88. [Google Scholar] [CrossRef]
- Ythier, E.; Lourenço, W.R. The geographical patterns of distribution of the genus Teuthraustes Simon, 1878 in Ecuador and description of three new species (Scorpiones, Chactidae); ZooKeys, 2017; pp. 45–63. [Google Scholar] [CrossRef] [PubMed]
- Lourenço, W.R. The third confirmed record of the scorpion genus Chactas Gervais, 1844 (Scorpiones, Chactidae) from Ecuador, with description of a new species from the Amazonian Province of Morona Santiago. Ent Mitt. Zool. MusHamburg 2014, 17, 171–8. [Google Scholar]
- Pocock, R.I. Descriptions of some new scorpions from Ecuador. Ann. Mag. Nat. Hist. 1898, 1, 413–422. [Google Scholar] [CrossRef]
- Botero-Trujillo, R.; Ochoa, J.A.; Prendini, L. A New Troglomorphic, Leaf-litter Scorpion from Ecuador (Troglotayosicidae: Troglotayosicus). Am. Mus. Novit. 2021, 2021, 1–24. [Google Scholar] [CrossRef]
- Sánchez-Vialas, A.; Blasco-Aróstegui, J.; García-Gila, J.; Lourenço, W.R. A new species of Troglotayosicus Lourenço, 1981 (Scorpiones: Troglotayosicidae) from southern Ecuador. Arachnology 2020, 18, 612–618. [Google Scholar] [CrossRef]
- Moreno-González, J.A.; Luna-Sarmiento, D.A.; Prendini, L. Phylogeny of the Troglomorphic Scorpion Genus Troglotayosicus (Scorpiones: Troglotayosicidae) with Description of a New Species from Colombia. Am. Mus. Novit. 2024, 2024, 1–39. [Google Scholar] [CrossRef]
- Botero-Trujillo, R.; González-Gómez, J.C.; Valenzuela-Rojas, J.C.; García, L.F. A new species in the troglomorphic scorpion genus Troglotayosicus from Colombia, representing the northernmost known record for the genus (Scorpiones, Troglotayosicidae). Zootaxa 2017, 4244, 568–582. [Google Scholar] [CrossRef] [PubMed]
- Lourenço, W.R.; Ythier, E. Description of Tityus (Atreus) cisandinus sp. n. from Ecuadorian Amazonia, with comments on some related species (Scorpiones: Buthidae). Arachn.-Riv. Aracnol. Ital. 2017, 15, 18. [Google Scholar]
- Kuntner, M. The seven grand challenges in arachnid science. Front Arachn. Sci. 2022, 1. [Google Scholar] [CrossRef]
- Štundlová, J.; Šťáhlavský, F.; Opatova, V.; Stundl, J.; Kovařík, F.; Dolejš, P.; et al. Molecular data do not support the traditional morphology-based groupings in the scorpion family Buthidae (Arachnida: Scorpiones). Mol. Phylogenet Evol. 2022, 173, 107511. [Google Scholar] [CrossRef] [PubMed]
- Santibáñez-López, C.E.; Ojanguren-Affilastro, A.A.; Graham, M.R.; Sharma, P.P. Congruence between ultraconserved element-based matrices and phylotranscriptomic datasets in the scorpion Tree of Life. Cladistics 2023, 39, 533–547. [Google Scholar] [CrossRef] [PubMed]
- Ballesteros, J.A.; Santibáñez-López, C.E.; Baker, C.M.; Benavides, L.R.; Cunha, T.J.; Gainett, G.; et al. Comprehensive Species Sampling and Sophisticated Algorithmic Approaches Refute the Monophyly of Arachnida. Mol. Biol. Evol. 2022, 39, msac021. [Google Scholar] [CrossRef] [PubMed]
- Sharma, P.P.; Fernández, R.; Esposito, L.A.; González-Santillán, E.; Monod, L. Phylogenomic resolution of scorpions reveals multilevel discordance with morphological phylogenetic signal. Proc. R Soc. B Biol. Sci. 2015, 282, 20142953. [Google Scholar] [CrossRef] [PubMed]
- Pocock, R.I. Descriptions of some new scorpions from Ecuador. Ann. Mag. Nat. Hist. 1898, 413–422. [Google Scholar] [CrossRef]
- Pocock, R.I. Some new or little-known Neotropical scorpions in the British Museum. J. Nat. Hist. 5, 469–478. [CrossRef]
- Lourenço, W.R.; Lourenço, W.R. La faune des Scorpions de l’Equateur. I. Les Buthidae. Systématique et biogéographie. Rev. Suisse Zool. 1988, 95, 681--697. [Google Scholar] [CrossRef]
- Thorell, T. Etudes scorpiologiques. Atti Della Soc. Ital. Sci. Nat. 1876, 19, 75–272. [Google Scholar]
- Simon, E. Études arachnologiques. 9e Mémoire. XV. Déscriptions de deux nouveaux genres de l’ordre des Scorpiones. Ann. Société Entomol. Fr. 1878, 5, 399–400. [Google Scholar]
- Borges, A.; Lomonte, B.; Angulo, Y.; Acosta de Patiño, H.; Pascale, J.M.; Otero, R.; et al. Venom diversity in the Neotropical scorpion genus Tityus: Implications for antivenom design emerging from molecular and immunochemical analyses across endemic areas of scorpionism. Acta Trop. 2020, 204, 105346. [Google Scholar] [CrossRef] [PubMed]
- Moreno-González, J.A.; Pinto-da-Rocha1, R.; Gallão2, J.E. Bringing order to a complex system: phenotypic and genotypic evidence contribute to the taxonomy of Tityus (Scorpiones, Buthidae) and support the description of a new species. ZooKeys 2021, 1075, 33–75. [Google Scholar] [CrossRef] [PubMed]
- Román, J.P.; García, F.; Medina, D.; Vásquez, M.; García, J.; Graham, M.R.; et al. Scorpion envenoming in Morona Santiago, Amazonian Ecuador: Molecular phylogenetics confirms involvement of the Tityus obscurus group. Acta Trop. 2018, 178, 1–9. [Google Scholar] [CrossRef] [PubMed]
- Wyman, J.T.; Wright-Ueda, J.; Agnew, Q.; Castellano, I.; Simone, Y. First report of arm-span competition in buthid scorpions: male-male contest in Tityus cf. rosenbergi Pocock, 1898. J. Arachnol. 2025, 52, 210–213. [Google Scholar] [CrossRef]
- Pardal, PP de O; Coelho, J.S.; da Silva, J.M.; Almeida, B.R.R.; Chalkidis, H.M.; Borges, A.; et al. Interpopulational genetic diversity in the medically important scorpion Tityus obscurus (Scorpiones: Buthidae) from northeastern Brazilian Amazonia. Acta Amaz. 2023, 53, 215–222. [Google Scholar] [CrossRef]
- Lourenço, W.R. Scorpion incidents, misidentification cases and possible implications for the final interpretation of results. J. Venom. Anim. Toxins Trop. Dis. 2016, 22, 21. [Google Scholar] [CrossRef] [PubMed]
- Blasco-Aróstegui, J.; Moreno-González, J.A.; Loria, S.F.; Carvalho, L.S.; Bird, T.L.; Nguyen, A.D.; et al. Include scorpions in global conservation plans. Science 2025, 389, 1099–1100. [Google Scholar] [CrossRef] [PubMed]
- Baradaran, M.; Salabi, F.; Mahdavinia, M.; Mohammadi, E.; Vazirianzadeh, B.; Avella, I.; et al. ScorpDb: A Novel Open-Access Database for Integrative Scorpion Toxinology. Toxins 2024, 16. [Google Scholar] [CrossRef] [PubMed]
- MSP. Gaceta Efectos Tóxicos–Ministerio de Salud Pública. 2025. Available online: https://www.salud.
- Wen, F.H.; Monteiro, W.M.; da Silva, A.M.M.; Tambourgi, D.V.; da Silva, I.M.; Sampaio, V.S.; et al. Snakebites and Scorpion Stings in the Brazilian Amazon: Identifying Research Priorities for a Largely Neglected Problem. PLoS Negl. Trop. Dis. 2015, 9, e0003701. [Google Scholar] [CrossRef] [PubMed]
- Guerra-Duarte, C.; Saavedra-Langer, R.; Matavel, A.; Oliveira-Mendes, B.B.R.; Chavez-Olortegui, C.; Paiva, A.L.B. Scorpion envenomation in Brazil: Current scenario and perspectives for containing an increasing health problem. PLoS Negl. Trop. Dis. 2023, 17, e0011069. [Google Scholar] [CrossRef] [PubMed]
- Borges, A.; Morales, M.; Loor, W.; Delgado, M. Scorpionism in Ecuador: First report of severe and fatal envenoming cases from northern Manabí by Tityus asthenes Pocock. Toxicon 2015, 105, 56–61. [Google Scholar] [CrossRef] [PubMed]
- Ochoa-Andrade, M.J.; Abril-López, P.A.; Molina Yanza, F.B.; Molina Gaibor, Á.A.; Guzmán León, J.L.; Niato Pacheco, J.; et al. Escorpionismo en la población amazónica del cantón Taisha en Ecuador. Rev Cubana Med Trop. 2022, 74. Available online: http://scielo.sld.cu/scielo.php?script=sci_abstract&pid=S0375-07602022000300013&lng=es&nrm=iso&tlng=es.
- Guerra-Duarte, C.; Saavedra-Langer, R.; Matavel, A.; Oliveira-Mendes, B.B.R.; Chavez-Olortegui, C.; Paiva, A.L.B. Scorpion envenomation in Brazil: Current scenario and perspectives for containing an increasing health problem. PLoS Negl. Trop. Dis. 2023, 17, e0011069. [Google Scholar] [CrossRef] [PubMed]
- Borges, A.; Román, J.P. Case Report: Fatal Scorpion Envenomation in a Shuar Child by Tityus cisandinus from Amazonian Ecuador: A Call for Specific Antivenom Availability in the Amazon Basin. Am. J. Trop. Med. Hyg. 2023, 108, 807–810. [Google Scholar] [CrossRef] [PubMed]
- Gaona Vásquez, O.A.; Quishpe Zagal, C.D.; Sotamba Quezada, J.R.; De La Cruz Novoa, M.G. ALACRANISMO EN EL PACIENTE PEDIÁTRICO: A PROPÓSITO DE UN CASO CLÍNICO. Rev. UNIANDES Cienc. Salud 2021, 4, 663–674. [Google Scholar]
- Khattabi, A.; Soulaymani-Bencheikh, R.; Achour, S.; Salmi, L.-R. for the Scorpion Consensus Expert Group. Classification of clinical consequences of scorpion stings: consensus development. Trans. R Soc. Trop. Med. Hyg. 2011, 105, 364–369. [Google Scholar] [CrossRef] [PubMed]
- Ministerio de Salud Pública. Manejo clínico de pacientes con mordeduras de serpientes venenosas y picaduras de escorpiones. Primera. Protocolo basado en la evidencia. In Primera; Dirección Nacional de Prevención y Control y Dirección de Normatización: Quito, 2017. [Google Scholar]
- Rodrigo, C.; Gnanathasan, A. Management of scorpion envenoming: a systematic review and meta-analysis of controlled clinical trials. Syst. Rev. 2017, 6, 74. [Google Scholar] [CrossRef] [PubMed]
- Vicepresidencia de la República del Ecuador. Vicepresidencia lideró coordinación interinstitucional que permitió entregar 40 dosis de suero antiescorpiónico para el país. Vicepresidencia [Internet]. Available online: https://www.vicepresidencia.gob.ec/vicepresidencia-lidero-coordinacion-interinstitucional-que-permitio-entregar-40-dosis-de-suero-antiescorpionico-para-el-pais/.
- Ratanabanangkoon: A simple and novel strategy for... - Google Scholar. 9 Apr 2026. Available online: https://journals.plos.org/plosntds/article?id=10.1371/journal.pntd.0004565.
- Bermúdez-Méndez, E.; Fuglsang-Madsen, A.; Føns, S.; Lomonte, B.; Gutiérrez, J.M.; Laustsen, A.H. Innovative Immunization Strategies for Antivenom Development. Toxins 2018, 10. [Google Scholar] [CrossRef] [PubMed]
- Darkaoui, B.; Hilal, I.; Khourcha, S.; Lafnoune, A.; Chakir, S.; Aarab, A.; et al. Development and Efficacy of the Antivenom Specific to Severe Envenomations in Morocco and North Africa: Advancements in Scorpion Envenomation Management. Toxins 2024, 16. [Google Scholar] [CrossRef] [PubMed]
- de Oliveira, I.S.; Alano-da-Silva, N.M.; Ferreira, I.G.; Cerni, F.A.; Sachett J de, A.G.; Monteiro, W.M.; et al. Understanding the complexity of Tityus serrulatus venom: A focus on high molecular weight components. J. Venom. Anim. Toxins Trop. Dis. 2024, 30, e20230046. [Google Scholar] [CrossRef] [PubMed]
- Costal-Oliveira, F.; Duarte, C.G.; Machado de Avila, R.A.; Melo, M.M.; Bordon, K.C.F.; Arantes, E.C.; et al. General biochemical and immunological characteristics of the venom from Peruvian scorpion Hadruroides lunatus. Toxicon 2012, 60, 934–942. [Google Scholar] [CrossRef] [PubMed]
- Borges, A.; Rojas de Arias, A.; Montaño, A.M.; de Souza, C.M.V. Scorpion Envenoming as an Emerging Public Health Problem in Paraguay, Bolivia, and Midwest Brazil: Involvement of Tityus confluens and the Need for a Panregional Evaluation of Available Antivenoms. Am. J. Trop. Med. Hyg. 2024, 111, 1166–1172. [Google Scholar] [CrossRef] [PubMed]
- Casewell, N.R.; Wüster, W.; Vonk, F.J.; Harrison, R.A.; Fry, B.G. Complex cocktails: the evolutionary novelty of venoms. Trends Ecol. Evol. 2013, 28, 219–229. [Google Scholar] [CrossRef] [PubMed]
- Herzig, V.; King, G.F.; Undheim, E.A.B. Can we resolve the taxonomic bias in spider venom research? Toxicon X 2019, 1, 100005. [Google Scholar] [CrossRef] [PubMed]
- Lüddecke, T.; Vilcinskas, A.; Lemke, S. Phylogeny-Guided Selection of Priority Groups for Venom Bioprospecting: Harvesting Toxin Sequences in Tarantulas as a Case Study. Toxins 2019, 11, 488. [Google Scholar] [CrossRef] [PubMed]
- von Reumont, B.M.; Campbell, L.I.; Jenner, R.A. Quo vadis venomics? A roadmap to neglected venomous invertebrates. Toxins 2014, 6, 3488–3551. [Google Scholar] [CrossRef] [PubMed]
- Abd El-Aziz, T.M.; Soares, A.G.; Stockand, J.D. Advances in venomics: Modern separation techniques and mass spectrometry. J. Chromatogr. B Anal. Technol. BioMed Life Sci. 2020, 1160, 122352. [Google Scholar] [CrossRef] [PubMed]
- Ducancel, F.; Durban, J.; Verdenaud, M. Transcriptomics and venomics: implications for medicinal chemistry. Future Med. Chem. 2014, 6, 1629–1643. [Google Scholar] [CrossRef] [PubMed]
- Lüddecke, T.; Paas, A.; Harris, R.J.; Talmann, L.; Kirchhoff, K.N.; Billion, A.; et al. Venom biotechnology: casting light on nature’s deadliest weapons using synthetic biology. Front Bioeng. Biotechnol. 2023, 11, 1166601. [Google Scholar] [CrossRef] [PubMed]
- Damm, M.; Vilcinskas, A.; Lüddecke, T. Mapping the architecture of animal toxin systems by mass spectrometry imaging. Biotechnol. Adv. 2025, 81, 108548. [Google Scholar] [CrossRef] [PubMed]
- Wiezel, G.A.; Bordon K de, C.F.; Martins, J.G.; Custódio VI do, C.; Matsuno, A.K.; Procópio, R.E.; de, L.; et al. A Hydrolase-Rich Venom Beyond Neurotoxins: Integrative Functional Proteomic and Immunoreactivity Analyses Reveal Novel Peptides in the Amazonian Scorpion Brotheas amazonicus. Int. J. Mol. Sci. 2026, 27, 1475. [Google Scholar] [CrossRef] [PubMed]
- Escobar, A.; Salazar, M.H.; Hernández-Ortiz, M.; Clement, H.; Encarnación-Guevara, S.; Cleghorn, J.; et al. Integration of proteomic and transcriptomic data of the venom and venom gland from Tityus jaimei. J. Proteom. 2026, 322, 105543. [Google Scholar] [CrossRef] [PubMed]
- Bordon, K.C.F.; Santos, G.C.; Martins, J.G.; Wiezel, G.A.; Amorim, F.G.; Crasset, T.; et al. Pioneering Comparative Proteomic and Enzymatic Profiling of Amazonian Scorpion Venoms Enables the Isolation of Their First α-Ktx, Metalloprotease, and Phospholipase A2. Toxins 2025, 17, 411. [Google Scholar] [CrossRef] [PubMed]
- Kalapothakis, Y.; Miranda, K.; Molina, D.A.M.; Conceição, I.M.C.A.; Larangote, D.; Op den Camp, H.J.M.; et al. An overview of Tityus cisandinus scorpion venom: Transcriptome and mass fingerprinting reveal conserved toxin homologs across the Amazon region and novel lipolytic components. Int. J. Biol. Macromol. 2023, 225, 1246–1266. [Google Scholar] [CrossRef] [PubMed]
- Delgado-Prudencio, G.; Cid-Uribe, J.I.; Morales, J.A.; Possani, L.D.; Ortiz, E.; Romero-Gutiérrez, T. The Enzymatic Core of Scorpion Venoms. Toxins 2022, 14, 248. [Google Scholar] [CrossRef] [PubMed]
- Lüddecke, T.; Hurka, S.; Dresler, J.; Lübcke, T.; von Wirth, V.; Lochnit, G.; et al. Comparative venomics suggests an evolutionary adaption of spider venom from predation to defense. Commun. Biol. 2025, 8, 1496. [Google Scholar] [CrossRef] [PubMed]
- Dresler, J.; Herzig, V.; Vilcinskas, A.; Lüddecke, T. Enlightening the toxinological dark matter of spider venom enzymes. npj Biodivers. 2024, 3, 25. [Google Scholar] [CrossRef] [PubMed]
- Dresler, J.; Avella, I.; Damm, M.; Dersch, L.; Krämer, J.; Vilcinskas, A.; et al. A roadmap to the enzymes from spider venom: biochemical ecology, molecular diversity, and value for the bioeconomy. Front Arachn. Sci. 2024, 3. [Google Scholar] [CrossRef]
- Calderón, J. Estudio transcriptómico de las glándulas de veneno de Otostigmus silvestrii (Myriapoda: Scolopendridae) y Teuthraustes atramentarius (Chelicerata: Chactidae): Estandarización de protocolos para especies de interés médico. Bachelor, Universidad San Francisco de Quito, 2024. [Google Scholar]
- Flores, E. Determination of the biological activities of the components present in the venom from the endemic Ecuadorian scorpion Teuthraustes aff. atramentarius; Bachelor, UNIVERSIDAD DE INVESTIGACIÓN DE TECNOLOGÍA EXPERIMENTAL YACHAY, 2020. [Google Scholar]
- Naseem, M.U.; Carcamo-Noriega, E.; Beltrán-Vidal, J.; Borrego, J.; Szanto, T.G.; Zamudio, F.Z.; et al. Cm28, a scorpion toxin having a unique primary structure, inhibits KV1.2 and KV1.3 with high affinity. J. Gen. Physiol. 2022, 154, e202213146. [Google Scholar] [CrossRef] [PubMed]
- Bartok, A.; Toth, A.; Somodi, S.; Szanto, T.G.; Hajdu, P.; Panyi, G.; et al. Margatoxin is a non-selective inhibitor of human Kv1.3 K+ channels. Toxicon 2014, 87, 6–16. [Google Scholar] [CrossRef] [PubMed]
- Garcia-Calvo, M.; Leonard, R.J.; Novick, J.; Stevens, S.P.; Schmalhofer, W.; Kaczorowski, G.J.; et al. Purification, characterization, and biosynthesis of margatoxin, a component of Centruroides margaritatus venom that selectively inhibits voltage-dependent potassium channels. J. Biol. Chem. 1993, 268, 18866–18874. [Google Scholar] [CrossRef]
- Lüddecke, T.; Herzig, V.; von Reumont, B.M.; Vilcinskas, A. The biology and evolution of spider venoms. Biol. Rev. Camb. Philos. Soc. 2022, 97, 163–178. [Google Scholar] [CrossRef] [PubMed]
- Ait Laaradia, M.; Laadraoui, J.; Ettitaou, A.; Agouram, F.; Oubella, K.; Moubtakir, S.; et al. Variation in venom yield, protein concentration and regeneration toxicity in the scorpion Buthus lienhardi. Toxicon Off. J. Int. Soc. Toxinology 2025, 255, 108254. [Google Scholar] [CrossRef] [PubMed]
- Agourram, Z.; Zegrari, R.; Kettani, A.; Badaoui, B.; Mkamel, M. Environmental determinants of venom variability in captive scorpions: A comprehensive analysis of diet, temperature, and humidity effects. Toxicon Off. J. Int. Soc. Toxinology 2024, 251, 108151. [Google Scholar] [CrossRef] [PubMed]
- Krämer, J.; Pommerening, R.; Predel, R. Equipped for Sexual Stings? Male-Specific Venom Peptides in Euscorpius italicus. Int. J. Mol. Sci. 2022, 23, 11020. [Google Scholar] [CrossRef] [PubMed]
- Reis, M.B.; Zoccal, K.F.; Gardinassi, L.G.; Faccioli, L.H. Scorpion envenomation and inflammation: Beyond neurotoxic effects. Toxicon 2019, 167, 174–179. [Google Scholar] [CrossRef] [PubMed]
- Kalapothakis, Y.; Miranda, K.; Molina, D.A.M.; Conceição, I.M.C.A.; Larangote, D.; Op den Camp, H.J.M.; et al. An overview of Tityus cisandinus scorpion venom: Transcriptome and mass fingerprinting reveal conserved toxin homologs across the Amazon region and novel lipolytic components. Int. J. Biol. Macromol. 2023, 225, 1246–1266. [Google Scholar] [CrossRef] [PubMed]
- Borges, A.; Morales, M.; Loor, W.; Delgado, M. Scorpionism in Ecuador: First report of severe and fatal envenoming cases from northern Manabí by Tityus asthenes Pocock. Toxicon 2015, 105, 56–61. [Google Scholar] [CrossRef] [PubMed]
| Family | Genus | Number of valid species | Region | Province | Natural History | Phenotype | ||
|---|---|---|---|---|---|---|---|---|
| Bothriuridae | Brachistoternus | 1 | Andes | Carchi | Fossorial | Medium size | ||
| Buthidae |
Ananteris | 3 |
Amazon, Andes, Coast | Azuay*, Cotopaxi*, Imbabura*, Los Rios, Manabí, Morona Santiago, Orellana*, Pichincha, Zamora Chinchipe* | Leaflitter |
Small size, brown-reddish spotted |
||
|
Centruroides |
3 | Coast, Insular |
Esmeraldas*, Galápagos, Guayas, Loja, Los Rios, Manabí, Santa Elena, | Often tree-dwelling Foliage-dwelling |
Large size, brown-yellow, sometimes dark, slender build, elongated metasoma in males | |||
| Tityus | 17 | Amazon, Andes, Coast | Carchi, Cotopaxi, El Oro, Esmeraldas, Imbabura, Loja, Manabi, Morona Santiago, Napo, Orellana, Pastaza, Pichincha, Santo Domingo de los Tsachilas, Sucumbios, Tungurahua, Zamora Chinchipe ^ |
Often tree-dwelling Foliage-dwelling |
Medium to large size, black to brown-reddish, elongated metasoma in males | |||
| Caraboctonidae | Hadruroides | 8 | Andes, Coast, Insular | Azuay, Bolivar, El Oro, Galapagos, Guayas, Loja, Manabi, Santa Elena |
Soil dwelling | Medium size, tones of yellow | ||
| Chactidae | Chactas | 3 | Amazon, Andes | Pichincha, Morona Santiago, Napo, Sucumbios |
Fossorial, Soil dwelling |
Small to medium size, brown-reddish | ||
| Teuthraustes | 15 | Amazon, Andes, Coast | Azuay, Bolivar, Cañar, Carchi¨, Chimborazo, Cotopaxi, El Oro, Esmeraldas, Guayas, Imbabura, Loja, Morona Santiago, Orellana, Pichincha, Santo Domingo de los Tsáchilas, Sucumbíos, Tungurahua, Zamora Chinchipe |
Fossorial, Soil dwelling | Medium size, black-brownish | |||
| Troglotayosicidae | Troglotayosicus | 3 | Amazon, Andes | Morona Santiago, Napo, Zamora Chinchipe | Troglobyotic, leaflitter | Small size, brown -reddish, absence of median eyes in the three species. |
||
| Year | 2016 | 2017 | 2018 | 2019 | 2020 | 2021 | 2022 | 2023 | 2024 | 2025 |
|---|---|---|---|---|---|---|---|---|---|---|
| Number of cases | 161 | 328 | 342 | 452 | 221 | 296 | 336 | 535 | 513 | 546 |
| Clinical feature/Reference | [69] | [60] | [73] | [72] |
| Number of cases | 5 cases (1, 2, 3, 6, and 16 years old) | 21 cases (1-40 years old) | 1 case (1 year old) | 1 case (4-month-old) |
| Province | Manabí | Morona Santiago | Orellana | Morona Santiago |
| Scorpion species | Tityus rosenbergi | Tityus cisandinus | Not determined | Tityus cisandinus |
| Sting location | Foot | Forearm, hand, gluteus, scrotum, thigh, leg, foot | Hand | Foot |
| Pharmacological treatment | saline solution, dexamethasone, acetaminophen, furosemide, ampicilline-sulbactam nebulized with ipratropium bromide and salbutamol, Oxigen, hydrocortisone, sodium penicillin G, ketorolac, clemastine, loratadine, metamizol metoclopramide |
Paracetamol, intravenous fluids, tramadol, diazepam, metoclopramide hydrochloride, potassium chloride. | Oxygen therapy (FiO₂), dextrose, midazolam, furosemide, Ringer’s lactate, insulin, benzodiazepines, normal saline, calcium gluconate, paracetamol, diazepam. | Alacramyn® scorpion antivenom (2 ampoules), dextrose, potassium chloride, ceftriaxone, hydrocortisone, tramadol, and calcium gluconate. |
| Fatalities | 1 (one year old) | 2 (four- and three-year-old) | No | 1 (4-month-old) |
| Class I: Local manifestations | ||||
| Pain | X | X | ||
| Erythema | X | X | X | |
| Edema | X | X | ||
| Bullous eruption | ||||
| Burning sensation | ||||
| Ecchymosis | ||||
| Hypereasthesia | ||||
| Itching | ||||
| Necrosis | ||||
| Paraesthesia | X | X | ||
| Purpura/Petechia | ||||
| Swelling Tingling | X | |||
| Class II: Minor manifestations (non-life threatening) | ||||
| Gastrointestinal effects | ||||
| Nausea/Vomiting (Emesis) | X | X | X | |
| Sialorrhea | X | X | X | |
| Abdominal distension | X | X | ||
| Nervous system effects | ||||
| Paleness | X | X | ||
| Sweating | X | X | X | |
| Clammy skin | X | X | ||
| Tachycardia | X | X | X | |
| Hypertension | ||||
| Hyperglycemia | X | X | X | |
| Hypotension | X | X | X | |
| Bradycardia | X | X | ||
| Miosis | X | |||
| Altered mental status | X | X | X | |
| Somnolence | X | X | X | X |
| Muscle weakness/rigidity | X | X | ||
| Cerebral decortication | X | |||
| Loss of corneal reflex | X | |||
| Seizures | X | |||
| Dysarthria | X | |||
| Headache | X | |||
| Muscle fasciculation | X | X | ||
| Mialgia | X | |||
| Respiratory effects | ||||
| Wheezing | X | |||
| Rales/bronchial breathing | X | X | X | |
| Reduced oxygen saturation | X | X | X | |
| Tachypnea | X | X | ||
| Dyspnea | X | X | ||
| Subcostal retractions | X | |||
| Cardiovascular effects | ||||
| Cardio-respiratory failure | X | X | X | |
| Myocarditis | ||||
| Hematological effects | ||||
| Leukocytosis | X | X | X | |
| Thrombocytosis | X | X | X | |
| Coagulopathy | X | |||
| Renal Effects | ||||
| Acute kidney injury | ||||
| General systemic effects | ||||
| Fever | X | X | ||
| Cyanosis | X | X |
| Family | Genus | Species | Proteome/Transcriptome | Toxins Venomzone |
Published Case Reports |
|---|---|---|---|---|---|
| Bothriuridae | Brachistosternus | pegnai | NO/NO | NO | NO |
| Buthidae | Ananteris | ashmolei | NO/NO | NO | NO |
| festae | NO/NO | NO | NO | ||
| mariaelenae | NO/NO | NO | NO | ||
| Centruroides | exsul | NO/NO | NO | NO | |
| gracilis* | NO/NO | SCX2_CENGR KGX52_CENGR KGX34_CENGR KGX13_CENGR |
NO | ||
| margaritatus* | NO/NO | KA321_CENMA KAX22_CENMA KAX48_CENMA KGX1A_CENMA SCX38_CENMA |
NO | ||
| Tityus | bastosi | NO/NO | NO | NO | |
| cisandinus | NO/YES [110] | No | YES [60,70,72] | ||
| crassicauda | NO/NO | NO | NO | ||
| demangei | NO/NO | NO | NO | ||
| ecuadorensis | NO/NO | NO | NO | ||
| forcipula | NO/NO | NO | NO | ||
| gasci | NO/NO | NO | NO | ||
| intermedius | NO/NO | NO | NO | ||
| julianae | NO/NO | NO | NO | ||
| jussarae | NO/NO | NO | NO | ||
| pugilator | NO/NO | NO | NO | ||
| roigi | NO/NO | NO | NO | ||
| rosenbergi | NO/NO | NO/NO | YES [111] | ||
| silvestris | NO/NO | NO | NO | ||
| simonsi | NO/NO | NO | NO | ||
| timendus | NO/NO | NO | NO | ||
| ythieri | NO/NO | NO | NO | ||
| Caraboctonidae | Hadruroides | charcasus | NO/NO | NO | NO |
| doriai | NO/NO | NO | NO | ||
| elenae | NO/NO | NO | NO | ||
| galapagoensis | NO/NO | NO | NO | ||
| maculatus | NO/NO | NO | NO | ||
| moreti | NO/NO | NO | NO | ||
| pachamama | NO/NO | NO | NO | ||
| udvardyi | NO/NO | NO | NO | ||
| Chactidae | Chactas | mahnerti | NO/NO | NO | NO |
| moreti | NO/NO | NO | NO | ||
| yaupi | NO/NO | NO | NO | ||
| Teuthraustes | atramentarius | NO/NO | NO | NO | |
| camposi | NO/NO | NO | NO | ||
| dubius | NO/NO | NO | NO | ||
| festae | NO/NO | NO | NO | ||
| gervaisii | NO/NO | NO | NO | ||
| giupponii | NO/NO | NO | NO | ||
| khodayarii | NO/NO | NO | NO | ||
| kuryi | NO/NO | NO | NO | ||
| lojanus | NO/NO | NO | NO | ||
| oculatus | NO/NO | NO | NO | ||
| ohausi | NO/NO | NO | NO | ||
| rosenbergi | NO/NO | NO | NO | ||
| simonsi | NO/NO | NO | NO | ||
| whymperi | NO/NO | NO | NO | ||
| wittii | NO/NO | NO | NO | ||
| Troglotayosicidae | Troglotayosicus | vachoni | NO/NO | NO | NO |
| muranunkae | NO/NO | NO | NO | ||
| ballvei | NO/NO | NO | NO |
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content. |
© 2026 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (http://creativecommons.org/licenses/by/4.0/).