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DOI10.47051/YKSW1188 Published January 17, 2021. Updated March 16, 2024. Open access.

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Common Mussurana (Clelia clelia)

Reptiles of Ecuador | Serpentes | Colubridae | Clelia clelia

English common names: Common Mussurana, Black Mussurana, Black Cribo.

Spanish common names: Ratonera común, chonta, lisa (Ecuador); cazadora negra (Colombia); zopilota, víbora de sangre (Juvenile), tiznada (Costa Rica); ratonera (Venezuela).

Recognition: ♂♂ 180 cmMaximum distance from the snout to the tip of the tail. ♀♀ 260 cmMaximum distance from the snout to the tip of the tail..¹ Clelia clelia is a large, robust snake with a cylindrical body and comparatively small eyes. The adults have a uniform glossy black or gray dorsum whereas young individuals measuring less than 60 cm in total length have a bright red dorsum with black scale tips, black head, and a cream or yellow nuchal collar followed by a black band (Fig. 1).^2,3 Individuals 60–90 cm in length are dull reddish brown to brownish black with a faint nape band.² The belly is always white but the dorsal coloration impinges the margins of the ventral scales.⁴ Clelia clelia is the only snake in Ecuador having this coloration and 19 rows of smooth scales at mid-body. This species differs from Drepanoides anomalus, Pseudoboa coronata, and C. equatoriana by having a greater number of dorsal scale rows at mid-body.^5–8

Figure 1: Individuals of Clelia clelia from Ecuador: Tiyu Yaku, Napo province (); Río Bigai Reserve, Orellana province (); La Selva Lodge, Sucumbíos province (); Zumba–Pucubamba road, Zamora Chinchipe province (). j=juvenile.

Natural history: Clelia clelia is a terrestrial and nocturnal snake that inhabits a variety of ecosystems ranging from savannas to cloud forests.^9–11 Mussuranas occur in old-growth forest, especially along bodies of water,^7,9 as well as in heavily disturbed areas such as pastures, cultivated fields,¹² yards, rural houses,¹³ and along roadsides.^9,14 Most individuals are seen active at night on the soil, leaf-litter, grass, swamps, creeks, or on rocky stream beds,^2,15 but in some cases, they have been spotted on tree branches up to 170 cm above the ground.^5,16 During the daytime, most individuals remain hidden, but others can be seen crossing roads and trails.^5,9 This snake feeds on other snakes, occasionally larger than itself.¹⁷ Prey include harmless snakes (Chironius fuscus, C. exoletus,¹⁸ Dipsas palmeri,¹⁹ Drymarchon corais,²⁰ Erythrolamprus reginae,² Helicops angulatus,²¹ Ninia hudsoni,²² Tantilla melanocephala, Xenodon rabdocephalus, X. severus,²³ as well as members of its own species), boas (Boa constrictor),^13,24 vipers (Bothriechis nigroadspersus,²⁵ Bothrops asper,²⁶ Lachesis muta, and Porthidium nasutum²⁷),²⁸ and coralsnakes (Micrurus obscurus).²³ Despite being primarily ophiophagous, Common Mussuranas also include in their diet: lizards (such as Ameiva ameiva,² Enyalioides heterolepis,⁹ and species of the genus Tupinambis), snake eggs, opossums,²⁹ rodents,⁵ birds, small mammals, and snails.^1,30 Individuals of C. clelia are active foragers, tracking prey by quickly flicking their tongues to detect their scent trail.⁹ The species is much appreciated by villagers because of its snake-eating habits.¹ Although it has grooved rear fangs and venom glands, this species also constricts its prey.^1,31 After striking it, the Mussurana subsequently launches the third of its body to surround the prey and then tightens it with body coils until the prey stops resisting.¹⁴ After that, it ingest the prey (dead or alive) head-first. This species has natural immunity to the venom of vipers.^32,33 Individuals are mostly calm. When grabbed, they constrict their bodies and do not usually strike, but instead hide their head under body coils and deploy a cloacal discharge.⁷ In humans, the venom of C. clelia can produce localized swelling, hemorrhage, and even necrosis (death of tissues and cells).³⁴

Females of Clelia clelia reach sexual maturity at 97.3 cm of snout-vent-length; males at 65 cm. During courtship, the female may respond aggressively at first and on occasion, can kill and eat the male.³¹ After a gestation period of 47 days,¹ females lay 9–25 eggs^1,14 that take 117–120 days to hatch.³⁵ In natural conditions, these are laid during periods of high rainfall. Hatchlings measure 31–49 cm in total length at birth.¹

Conservation: Least Concern Believed to be safe from extinction given current circumstances..³⁶ Clelia clelia is listed in this category mainly on the basis of the species’ wide distribution, occurrence in protected areas, presumed stable populations, and adaptability to human-modified environments.³⁶ However, the decline in the abundance of prey, coupled with the destruction and fragmentation of forested environments throughout Central and South America can be a threat for the long-term of survival of the species.³⁷ In a rainforest locality in Panamá, the occurrence rates of C. clelia have diminished in the period from 1997 to 2012,³⁸ probably as a result of the diminished snake abundance caused by a corresponding loss of amphibians.³⁸ Additionally, individuals of C. clelia are commonly found dead on the roads due vehicular traffic.³⁹

Distribution: Clelia clelia is widely-distributed in Central and South America, from southern México (Yucatán) to northern Argentina.^11,36 The species has an estimated total range size of 2,078,373 km² that encompasses much of Mesoamerica, the Chocó, Magdalena valley, the Llanos plains, the Amazon rainforest, and El Chaco.¹¹ In Ecuador, this species occurs at both sides of the Andes (Fig. 2).

Figure 2: Distribution of Clelia clelia in Ecuador. See Appendix 1 for a complete list of the presence localities included in the map.

Etymology: The word clelia is derived from the Latin Cloelia, a girl’s name meaning “illustrious” or “famous.” According to Roman legend, Cloelia was a heroine who was held hostage by an Etruscan invader. However, she managed to escape by swimming across the river Tiber.⁴⁰

See it in the wild: Some of the best localities to find Common Mussuranas in the wild in Ecuador are Yasuní Scientific Station, Jatun Sacha Biological Reserve, and Jama Coaque Reserve. The snakes are more easily located by walking along forested rivers and streams at night.

Acknowledgments: Special thanks to Darwin Núñez and Diego Piñán for providing locality data for Clelia clelia.

Authors: Juan C. Díaz-Ricaurte^aAffiliation: Universidade de São Paulo, São Paulo, Brazil. and Alejandro Arteaga^bAffiliation: Tropical Herping (TH), Quito, Ecuador.

Photographers: Jose Vieira^bAffiliation: Tropical Herping (TH), Quito, Ecuador.^,cAffiliation: ExSitu, Quito, Ecuador. and Sebastián Di Doménico^dAffiliation: Keeping Nature, Bogotá, Colombia.

How to cite? Díaz-Ricaurte JC, Arteaga A (2024) Common Mussurana (Clelia clelia). In: Arteaga A, Bustamante L, Vieira J (Eds) Reptiles of Ecuador: Life in the middle of the world. Available from: www.reptilesofecuador.com. DOI: 10.47051/YKSW1188

Literature cited:

1. Savage JM (2002) The amphibians and reptiles of Costa Rica, a herpetofauna between two continents, between two seas. The University of Chicago Press, Chicago, 934 pp.
2. Duellman WE (1978) The biology of an equatorial herpetofauna in Amazonian Ecuador. Publications of the Museum of Natural History, University of Kansas 65: 1–352.
3. Arquilla AM, Lehtinen RM (2018) Geographic variation in head band shape in juveniles of Clelia clelia (Colubridae). Mesoamerican Herpetology 5: 112–120.
4. Murphy JC, Downie R, Smith JM, Livingstone S, Mohammed R, Lehtinen RM, Eyre M, Sewlal JN, Noriega N, Casper GS, Anton T, Rutherford MG, Braswell AL, Jowers MJ (2018) A field guide to the amphibians & reptiles of Trinidad and Tobago. Trinidad and Tobago Naturalist’s Club, Port of Spain, 336 pp.
5. Duellman WE (2005) Cusco amazónico: the lives of amphibians and reptiles in an Amazonian rainforest. Cornell University Press, Ithaca, 433 pp.
6. de Fraga R, Lima AP, da Costa Prudente AL, Magnusson WE (2013) Guia de cobras da região de Manaus - Amazônia Central. Editopa Inpa, Manaus, 303 pp.
7. Martins M, Oliveira ME (1998) Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil. Herpetological Natural History 6: 78–150.
8. Arteaga A, Bustamante L, Guayasamin JM (2013) The amphibians and reptiles of Mindo. Universidad Tecnológica Indoamérica, Quito, 257 pp.
9. Field notes, Reptiles of Ecuador book project.
10. França FGR, Mesquita DO, Colli GR (2006) A checklist of snakes from Amazonian savannas in Brazil, housed in the Coleção Herpetológica da Universidade de Brasília, with new distribution records. Occasional Papers of the Sam Noble Oklahoma Museum of Natural History 17: 1–13.
11. Nogueira CC, Argôlo AJS, Arzamendia V, Azevedo JA, Barbo FE, Bérnils RS, Bolochio BE, Borges-Martins M, Brasil-Godinho M, Braz H, Buononato MA, Cisneros-Heredia DF, Colli GR, Costa HC, Franco FL, Giraudo A, Gonzalez RC, Guedes T, Hoogmoed MS, Marques OAV, Montingelli GG, Passos P, Prudente ALC, Rivas GA, Sanchez PM, Serrano FC, Silva NJ, Strüssmann C, Vieira-Alencar JPS, Zaher H, Sawaya RJ, 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. DOI: 10.2994/SAJH-D-19-00120.1
12. Lynch JD (2015) The role of plantations of the African palm (Elaeis guineensis Jacq.) in the conservation of snakes in Colombia. Caldasia 37: 169–182.
13. Beebe W (1946) Field notes on the snakes of Kartabo, British Guiana, and Caripito, Venezuela. Zoologica 31: 11–52.
14. Leenders T (2019) Reptiles of Costa Rica: a field guide. Cornell University Press, Ithaca, 625 pp.
15. Solórzano A (2004) Serpientes de Costa Rica. Distribución, taxonomía e historia natural. Universidad de Costa Rica, San José, 792 pp.
16. Photographic record by César Barrio-Amorós.
17. César Barrio-Amorós, pers. comm.
18. Photo by Patrick Campbell.
19. MECN, JOCOTOCO, ECOMINGA (2013) Herpetofauna en áreas prioritarias para la conservación: el sistema de reservas Jocotoco y Ecominga. Museo Ecuatoriano de Ciencias Naturales, Quito, 408 pp.
20. Photo by Giuseppe Gagliardi-Urrutia.
21. Photo by Vincent Vos.
22. Wright T, Floyd E, Camper JD, Nilsson J (2019) Clelia clelia (Black Mussurana). Diet. Herpetological Review 50: 388–387.
23. Harry Turner, pers. comm.
24. Photo by Josua Hannink.
25. Chavarría M, Barrio-Amorós C (2014) Clelia clelia. Predation. Mesoamerican Herpetology 1: 286.
26. Campbell JA, Lamar WW (2004) The venomous reptiles of the western hemisphere. Cornell University Press, Ithaca, 774 pp.
27. Delia J (2009) Another crotaline snake prey item of the Neotropical snake Clelia clelia (Daudin 1983). Herpetology Notes 2: 21–22.
28. Valencia JH, Garzón-Tello K, Barragán-Paladines ME (2016) Serpientes venenosas del Ecuador: sistemática, taxonomía, historial natural, conservación, envenenamiento y aspectos antropológicos. Fundación Herpetológica Gustavo Orcés, Quito, 653 pp.
29. Martins Teixeira D, Lorini ML, Persson VG, Porto M (1991) Clelia clelia (Musurana). Feeding behavior. Herpetological Review 22: 131–132.
30. Alencar LR, Gaiarsa MP, Martins M (2013) The evolution of diet and microhabitat use in Pseudoboine Snakes. South American Journal of Herpetology 8: 60–66. DOI: 10.2994/sajh-d-13-00005.1
31. Scott Jr NJ (1983) Clelia clelia. In: Janzen DH (Ed) Costa Rican natural history. The University of Chicago Press, Chicago, 392–393.
32. Cerdas L, Lomonte B (1982) Estudio de la capacidad ofiófaga y la resistencia de la zopilota (Clelia clelia, Colubridae) de Costa Rica a los venenos de serpiente. Toxicon 20: 936–939. DOI: 10.1016/0041-0101(82)90083-6
33. Lomonte B, Cerdas L, Solórzano A, Martínez S (1989) El suero de neonatos de Clelia clelia (Serpentes: Colubridae) neutraliza la acción hemorrágica del veneno de Bothrops asper (Serpentes: Viperidae). Revista de Biología Tropical 38: 325–326.
34. Weinstein SA, Warrell DA, White J, Keyler DE (2011) “Venomous” bites from non-venomous snakes. Elsevier, Amsterdam, 364 pp. DOI: 10.1016/C2010-0-68461-6
35. Martínez S, Cerdas-Fallas L (1986) Captive reproduction of the Mussurana, Clelia clelia Daudin from Costa Rica. Herpetological Review 17: 12.
36. Gutiérrez-Cárdenas P, Rivas G, Nogueira C, Gagliardi G, Catenazzi A, Gonzáles L, Murphy J (2019) Clelia clelia. The IUCN Red List of threatened species. Available from: www.iucnredlist.org. DOI: 10.2305/IUCN.UK.2019-3.RLTS.T197468A2487325.en
37. Lynch JD (2012) El contexto de las serpientes de Colombia con un análisis de las amenazas contra su conservación. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 36: 435–449.
38. Zipkin EF, DiRenzo GD, Ray JM, Rossman S, Lips KR (2020) Tropical snake diversity collapses after widespread amphibian loss. Science 367: 814–816. DOI: 10.1126/science.aay5733
39. Vargas-Salinas F, Delgado-Ospina I, López-Aranda F (2011) Mortalidad por atropello vehicular y distribución de anfibios y reptiles en un bosque subandino en el occidente de Colombia. Caldasia 33: 121–138.
40. Beolens B, Watkins M, Grayson M (2011) The eponym dictionary of reptiles. Johns Hopkins University Press, Baltimore, 296 pp.

Appendix 1: Locality data used to create the distribution map of Clelia clelia in Ecuador (Fig. 2). Go to the section on symbols and abbreviations for a list of acronyms used.

Country	Province	Locality	Source
Colombia	Caquetá	Belén de Andaquíes	Díaz-Ricaurte et al. 2018
Colombia	Caquetá	Caserío La Rastra	Díaz-Ricaurte et al. 2018
Colombia	Caquetá	Florencia	Díaz-Ricaurte et al. 2018
Colombia	Caquetá	Morelia	iNaturalist; photo examined
Colombia	Caquetá	Santuarios	Ruiz Valderrama 2023
Colombia	Caquetá	Vereda Fátima	Cárdenas Hincapié & Lozano Bernal 2023
Colombia	Caquetá	Via antigua Caquetá–Huila	Díaz-Ricaurte et al. 2018
Colombia	Nariño	Boca del Río Mira	Carvajal et al. 2024
Colombia	Nariño	Cabo Manglares	Higuera Rojas et al. 2021
Colombia	Putumayo	La Paya	Peña Alzáte et al. 2020
Colombia	Putumayo	Piñuña, 22 km NE of	Hoyos et al. 2023
Colombia	Putumayo	Puerto Asís	Díaz-Ricaurte et al. 2018
Colombia	Putumayo	Puerto Leguizamo, 25 km N of	Parques Nacionales de Colombia
Ecuador	Cañar	Terminal La Troncal	Ortega Torres 2015
Ecuador	El Oro	Machala	Nogueira et al. 2019
Ecuador	Esmeraldas	Atacames	Nogueira et al. 2019
Ecuador	Esmeraldas	Bilsa Biological Reserve	Reptiles of Ecuador book database
Ecuador	Esmeraldas	Esmeraldas	Zaher 1996
Ecuador	Esmeraldas	Pajonal	Morales 2004
Ecuador	Esmeraldas	Palmicultora La Tolita	Online multimedia
Ecuador	Esmeraldas	San Lorenzo	Zaher 1996
Ecuador	Guayas	Balzar	Zaher 1996
Ecuador	Guayas	Guayaquil	Zaher 1996
Ecuador	Guayas	Río Daule	MCZ 3570; VertNet
Ecuador	Loja	Bosque Petrificado Puyango	Garzón-Santomaro et al. 2019
Ecuador	Loja	Loja	Nogueira et al. 2019
Ecuador	Loja	Yangana	Arteaga et al. 2013
Ecuador	Los Ríos	Centro Científico Río Palenque	MCZ 152749; VertNet
Ecuador	Manabí	Bosque Seco Lalo Loor	Reptiles of Ecuador book database
Ecuador	Manabí	El Guayacán	iNaturalist; photo examined
Ecuador	Manabí	Jama Coaque Reserve	iNaturalist; photo examined
Ecuador	Manabí	La Unión de Santa Ana	MHNG 2531.061; collection database
Ecuador	Manabí	Maicito	MHNG 1357.014; collection database
Ecuador	Manabí	Pedernales, 15 km SE of	Pazmiño-Otamendi 2020
Ecuador	Manabí	Río Jama	Photo by David Salazar
Ecuador	Manabí	San Isidro	Cisneros-Heredia et al. 2007
Ecuador	Morona Santiago	Bosque Protector Abanico	Cumba Endara 2008
Ecuador	Morona Santiago	Macas	Zaher 1996
Ecuador	Morona Santiago	Macas, 14 km W of	Nogueira et al. 2019
Ecuador	Morona Santiago	Macuma	Zaher 1996
Ecuador	Morona Santiago	Méndez–El Pescado road	Valencia et al. 2016
Ecuador	Morona Santiago	Normandía	Zaher 1996
Ecuador	Morona Santiago	Quebrada Napinaza	Nogueira et al. 2019
Ecuador	Morona Santiago	Reserva Ecológica El Paraíso	Photo by Alex Achig
Ecuador	Morona Santiago	Río Santiago	Zaher 1996
Ecuador	Morona Santiago	Río Upano	iNaturalist; photo examined
Ecuador	Morona Santiago	Santa Ana, 1 km NE of	Reptiles of Ecuador book database
Ecuador	Morona Santiago	Sardinayacu	Nogueira et al. 2019
Ecuador	Morona Santiago	Taisha	Valencia et al. 2016
Ecuador	Morona Santiago	Vall del Río Quimi	Betancourt et al. 2018
Ecuador	Napo	Archidona	Nogueira et al. 2019
Ecuador	Napo	Archidona, 1.5 km N of	iNaturalist; photo examined
Ecuador	Napo	Baeza	Nogueira et al. 2019
Ecuador	Napo	El Chaco	Nogueira et al. 2019
Ecuador	Napo	El Chaco, 1 km NE of	Zaher 1996
Ecuador	Napo	Finca Fischer	TCWC 65016; VertNet
Ecuador	Napo	Gareno Lodge	Photo by Margy Green
Ecuador	Napo	Guagua Sumaco	iNaturalist; photo examined
Ecuador	Napo	Guamaní	Nogueira et al. 2019
Ecuador	Napo	Hidroeléctrica Coca Codo Sinclair	MECN & ENTRIX 2009–2013
Ecuador	Napo	Ikiam	Reptiles of Ecuador book database
Ecuador	Napo	Jatun Sacha Biological Station	MCZ 173840; VertNet
Ecuador	Napo	Limoncocha	UIMNH 54662; VertNet
Ecuador	Napo	Misahuallí	Reptiles of Ecuador book database
Ecuador	Napo	Río Bigai Reserve	Reptiles of Ecuador book database
Ecuador	Napo	Rio Hollín	Nogueira et al. 2019
Ecuador	Napo	Río Huataraco	Nogueira et al. 2019
Ecuador	Napo	Río Oyacachi	Zaher 1996
Ecuador	Napo	Río Quijos Ecolodge	Reptiles of Ecuador book database
Ecuador	Napo	Río Suno	Zaher 1996
Ecuador	Napo	Runa Huasi	Nogueira et al. 2019
Ecuador	Napo	San Francisco de Borja	USNM 210855; VertNet
Ecuador	Napo	Tena	iNaturalist; photo examined
Ecuador	Napo	Tena, 4 km N of	iNaturalist; photo examined
Ecuador	Napo	Tiyu Yaku	Reptiles of Ecuador book database
Ecuador	Napo	Wildsumaco Wildlife Sanctuary	Camper et al. 2021
Ecuador	Napo	Yachana Reserve	Whitworth & Beirne 2011
Ecuador	Napo	Zatzayacu, 4.5 km NNE	KU 146735; VertNet
Ecuador	Napo	Zoo el Arca	Photo by Diego Piñán
Ecuador	Orellana	Ávila Viejo	Nogueira et al. 2019
Ecuador	Orellana	Campo Apaika	Nogueira et al. 2019
Ecuador	Orellana	Cordillera Galeras	Nogueira et al. 2019
Ecuador	Orellana	Dicaro, 1.8 km NW of	Photo by Morley Read
Ecuador	Orellana	El Coca	Nogueira et al. 2019
Ecuador	Orellana	Estación Científica Yasuní	Nogueira et al. 2019
Ecuador	Orellana	Loreto	Reptiles of Ecuador book database
Ecuador	Orellana	NPF, 5 km N of	Nogueira et al. 2019
Ecuador	Orellana	Payamino, 6 km SW of	iNaturalist; photo examined
Ecuador	Orellana	Reserva Río Bigal	García et al. 2021
Ecuador	Orellana	Río Payamino	iNaturalist; photo examined
Ecuador	Orellana	San José de Payamino	iNaturalist; photo examined
Ecuador	Orellana	Shiripuno Lodge	Online multimedia
Ecuador	Orellana	SPF, 12 km N of	Nogueira et al. 2019
Ecuador	Orellana	Vía Maxus, km 98	Nogueira et al. 2019
Ecuador	Pastaza	Alto Curaray	USNM 210852
Ecuador	Pastaza	Alto Río Curaray	Zaher 1996
Ecuador	Pastaza	Balsaura	Ortega-Andrade 2010
Ecuador	Pastaza	Conambo	Ortega-Andrade 2010
Ecuador	Pastaza	Juyuintza	Ortega-Andrade 2010
Ecuador	Pastaza	Kapawi Ecolodge	Reptiles of Ecuador book database
Ecuador	Pastaza	Lago Gawope	Nogueira et al. 2019
Ecuador	Pastaza	Mera	Nogueira et al. 2019
Ecuador	Pastaza	Mera, 3 km N of	iNaturalist; photo examined
Ecuador	Pastaza	Pindoyacu	Ortega-Andrade 2010
Ecuador	Pastaza	Río Anzu Reserve	Reyes-Puig et al. 2013
Ecuador	Pastaza	Río Bufeo	Ortega-Andrade 2010
Ecuador	Pastaza	Río Lliquino	Zaher 1996
Ecuador	Pastaza	Río Punino	Nogueira et al. 2019
Ecuador	Pastaza	Río Villano	Nogueira et al. 2019
Ecuador	Pastaza	San José de Curaray	iNaturalist; photo examined
Ecuador	Pastaza	Santa Ana	Nogueira et al. 2019
Ecuador	Pastaza	Santa Clara	Nogueira et al. 2019
Ecuador	Pastaza	Villano	Reptiles of Ecuador book database
Ecuador	Sucumbíos	Dureno	Yánez-Muñoz & Chimbo 2007
Ecuador	Sucumbíos	El Eno, 4.7 km N of	Nogueira et al. 2019
Ecuador	Sucumbíos	El Reventador	MHNG 2445.005
Ecuador	Sucumbíos	Estación Amazonas OCP	Valencia & Garzón 2011
Ecuador	Sucumbíos	Estación Cayagama	Valencia & Garzón 2011
Ecuador	Sucumbíos	Estación PUCE en Cuyabeno	Nogueira et al. 2019
Ecuador	Sucumbíos	Garzacocha	iNaturalist; photo examined
Ecuador	Sucumbíos	La Selva Lodge	Reptiles of Ecuador book database
Ecuador	Sucumbíos	Lagartococha	iNaturalist; photo examined
Ecuador	Sucumbíos	Laguna Grande	Nogueira et al. 2019
Ecuador	Sucumbíos	Limoncocha	LACM 73327; VertNet
Ecuador	Sucumbíos	Mushullacta	Nogueira et al. 2019
Ecuador	Sucumbíos	Pisorié	Yánez-Muñoz & Chimbo 2007
Ecuador	Sucumbíos	Pozo Lago Norte	iNaturalist; photo examined
Ecuador	Sucumbíos	Río Malo	Nogueira et al. 2019
Ecuador	Sucumbíos	San Pablo de Kantesiya	Nogueira et al. 2019
Ecuador	Sucumbíos	Sani Lodge	Reptiles of Ecuador book database
Ecuador	Sucumbíos	Santa Cecilia	Nogueira et al. 2019
Ecuador	Sucumbíos	Tarapoa	Nogueira et al. 2019
Ecuador	Tungurahua	Cerro Candelaria Reserve	Reptiles of Ecuador book database
Ecuador	Tungurahua	Reserva Río Zuñac	Yánez-Muñoz et al. 2013
Ecuador	Zamora Chinchipe	Alto Machinaza	Nogueira et al. 2019
Ecuador	Zamora Chinchipe	Cerro Plateado	Darwin Núñez, pers. comm.
Ecuador	Zamora Chinchipe	Ciudad Perdida	Online multimedia
Ecuador	Zamora Chinchipe	El Chorro, 1.4 km S of	Reptiles of Ecuador book database
Ecuador	Zamora Chinchipe	Jamboé	Darwin Núñez, pers. comm.
Ecuador	Zamora Chinchipe	Jambué Bajo	Nogueira et al. 2019
Ecuador	Zamora Chinchipe	Loja, 21 km E of	Zaher 1996
Ecuador	Zamora Chinchipe	Reserva Biológica Cerro Plateado	Nogueira et al. 2019
Ecuador	Zamora Chinchipe	Via Romerillos	Darwin Núñez, pers. comm.
Ecuador	Zamora Chinchipe	Zamora	Cisneros-Heredia et al. 2007
Ecuador	Zamora Chinchipe	Zumba–Pucubamba road	This work; Fig. 1
Perú	Amazonas	Chiriaco, 43 km NE of	LSUMZ 39325; VertNet
Perú	Amazonas	Cumba	Koch et al. 2018
Perú	Amazonas	Quebrada Honda	Koch et al. 2018
Perú	Cajamarca	Perico	Koch et al. 2018
Perú	Loreto	Parinari	Nogueira et al. 2019

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