Published January 17, 2021. Updated March 16, 2024. Open access.

Gallery ❯

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.58

Figure showing variation among individuals of Clelia clelia

Figure 1: Individuals of Clelia clelia from Ecuador: Tiyu Yaku, Napo province (); Reserva Río Bigai, 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.911 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

“The bright red juveniles of the two Costa Rican mussuranas are considered to be víboras de sangre (blood snakes) whose bite results in bleeding over the entire body surface and death. This in an unfounded belief lacking any element of fact.”

Jay M Savage, American herpetologist, 2002.

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 eggs1,14 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

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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 Panama, 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 Mexico (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).

Distribution of Clelia clelia in Ecuador

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-RicaurteaAffiliation: Universidade de São Paulo, São Paulo, Brazil. and Alejandro ArteagabAffiliation: Tropical Herping (TH), Quito, Ecuador.

Photographers: Jose VieirabAffiliation: Tropical Herping (TH), Quito, Ecuador.,cAffiliation: ExSitu, Quito, Ecuador. and Sebastián Di DoménicodAffiliation: 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: 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: 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.

ColombiaCaquetáBelén de AndaquíesDíaz-Ricaurte et al. 2018
ColombiaCaquetáCaserío La RastraDíaz-Ricaurte et al. 2018
ColombiaCaquetáFlorenciaDíaz-Ricaurte et al. 2018
ColombiaCaquetáMoreliaiNaturalist; photo examined
ColombiaCaquetáSantuariosRuiz Valderrama 2023
ColombiaCaquetáVereda FátimaCárdenas Hincapié & Lozano Bernal 2023
ColombiaCaquetáVia antigua Caquetá–HuilaDíaz-Ricaurte et al. 2018
ColombiaNariñoBoca del Río MiraCarvajal et al. 2024
ColombiaNariñoCabo ManglaresHiguera Rojas et al. 2021
ColombiaPutumayoLa PayaPeña Alzáte et al. 2020
ColombiaPutumayoPiñuña, 22 km NE ofHoyos et al. 2023
ColombiaPutumayoPuerto AsísDíaz-Ricaurte et al. 2018
ColombiaPutumayoPuerto Leguizamo, 25 km N ofParques Nacionales de Colombia
EcuadorCañarTerminal La TroncalOrtega Torres 2015
EcuadorEl OroMachalaNogueira et al. 2019
EcuadorEsmeraldasAtacamesNogueira et al. 2019
EcuadorEsmeraldasBilsa Biological ReserveReptiles of Ecuador book database
EcuadorEsmeraldasEsmeraldasZaher 1996
EcuadorEsmeraldasPajonalMorales 2004
EcuadorEsmeraldasPalmicultora La TolitaOnline multimedia
EcuadorEsmeraldasSan LorenzoZaher 1996
EcuadorGuayasBalzarZaher 1996
EcuadorGuayasGuayaquilZaher 1996
EcuadorGuayasRío DauleMCZ 3570; VertNet
EcuadorLojaBosque Petrificado PuyangoGarzón-Santomaro et al. 2019
EcuadorLojaLojaNogueira et al. 2019
EcuadorLojaYanganaArteaga et al. 2013
EcuadorLos RíosCentro Científico Río PalenqueMCZ 152749; VertNet
EcuadorManabíBosque Seco Lalo LoorReptiles of Ecuador book database
EcuadorManabíEl GuayacániNaturalist; photo examined
EcuadorManabíJama Coaque ReserveiNaturalist; photo examined
EcuadorManabíLa Unión de Santa AnaMHNG 2531.061; collection database
EcuadorManabíMaicitoMHNG 1357.014; collection database
EcuadorManabíPedernales, 15 km SE ofPazmiño-Otamendi 2020
EcuadorManabíRío JamaPhoto by David Salazar
EcuadorManabíSan IsidroCisneros-Heredia et al. 2007
EcuadorMorona SantiagoBosque Protector AbanicoCumba Endara 2008
EcuadorMorona SantiagoMacasZaher 1996
EcuadorMorona SantiagoMacas, 14 km W ofNogueira et al. 2019
EcuadorMorona SantiagoMacumaZaher 1996
EcuadorMorona SantiagoMéndez–El Pescado roadValencia et al. 2016
EcuadorMorona SantiagoNormandíaZaher 1996
EcuadorMorona SantiagoQuebrada NapinazaNogueira et al. 2019
EcuadorMorona SantiagoReserva Ecológica El Paraíso Photo by Alex Achig
EcuadorMorona SantiagoRío SantiagoZaher 1996
EcuadorMorona SantiagoRío UpanoiNaturalist; photo examined
EcuadorMorona SantiagoSanta Ana, 1 km NE ofReptiles of Ecuador book database
EcuadorMorona SantiagoSardinayacuNogueira et al. 2019
EcuadorMorona SantiagoTaishaValencia et al. 2016
EcuadorMorona SantiagoVall del Río QuimiBetancourt et al. 2018
EcuadorNapoArchidonaNogueira et al. 2019
EcuadorNapoArchidona, 1.5 km N ofiNaturalist; photo examined
EcuadorNapoBaezaNogueira et al. 2019
EcuadorNapoEl ChacoNogueira et al. 2019
EcuadorNapoEl Chaco, 1 km NE ofZaher 1996
EcuadorNapoFinca FischerTCWC 65016; VertNet
EcuadorNapoGareno LodgePhoto by Margy Green
EcuadorNapoGuagua SumacoiNaturalist; photo examined
EcuadorNapoGuamaníNogueira et al. 2019
EcuadorNapoHidroeléctrica Coca Codo SinclairMECN & ENTRIX 2009–2013
EcuadorNapoIkiamReptiles of Ecuador book database
EcuadorNapoJatun Sacha Biological StationMCZ 173840; VertNet
EcuadorNapoLimoncochaUIMNH 54662; VertNet
EcuadorNapoMisahuallíReptiles of Ecuador book database
EcuadorNapoReserva Río BigaiReptiles of Ecuador book database
EcuadorNapoRio HollínNogueira et al. 2019
EcuadorNapoRío HuataracoNogueira et al. 2019
EcuadorNapoRío OyacachiZaher 1996
EcuadorNapoRío Quijos EcolodgeReptiles of Ecuador book database
EcuadorNapoRío SunoZaher 1996
EcuadorNapoRuna HuasiNogueira et al. 2019
EcuadorNapoSan Francisco de BorjaUSNM 210855; VertNet
EcuadorNapoTenaiNaturalist; photo examined
EcuadorNapoTena, 4 km N ofiNaturalist; photo examined
EcuadorNapoTiyu YakuReptiles of Ecuador book database
EcuadorNapoWildsumaco Wildlife SanctuaryCamper et al. 2021
EcuadorNapoYachana ReserveWhitworth & Beirne 2011
EcuadorNapoZatzayacu, 4.5 km NNE KU 146735; VertNet
EcuadorNapoZoo el ArcaPhoto by Diego Piñán
EcuadorOrellanaÁvila ViejoNogueira et al. 2019
EcuadorOrellanaCampo ApaikaNogueira et al. 2019
EcuadorOrellanaCordillera GalerasNogueira et al. 2019
EcuadorOrellanaDicaro, 1.8 km NW ofPhoto by Morley Read
EcuadorOrellanaEl CocaNogueira et al. 2019
EcuadorOrellanaEstación Científica YasuníNogueira et al. 2019
EcuadorOrellanaLoretoReptiles of Ecuador book database
EcuadorOrellanaNPF, 5 km N ofNogueira et al. 2019
EcuadorOrellanaPayamino, 6 km SW ofiNaturalist; photo examined
EcuadorOrellanaReserva Río BigalGarcía et al. 2021
EcuadorOrellanaRío PayaminoiNaturalist; photo examined
EcuadorOrellanaSan José de PayaminoiNaturalist; photo examined
EcuadorOrellanaShiripuno LodgeOnline multimedia
EcuadorOrellanaSPF, 12 km N ofNogueira et al. 2019
EcuadorOrellanaVía Maxus, km 98Nogueira et al. 2019
EcuadorPastazaAlto CurarayUSNM 210852
EcuadorPastazaAlto Río CurarayZaher 1996
EcuadorPastazaBalsauraOrtega-Andrade 2010
EcuadorPastazaConamboOrtega-Andrade 2010
EcuadorPastazaJuyuintzaOrtega-Andrade 2010
EcuadorPastazaKapawi EcolodgeReptiles of Ecuador book database
EcuadorPastazaLago GawopeNogueira et al. 2019
EcuadorPastazaMeraNogueira et al. 2019
EcuadorPastazaMera, 3 km N ofiNaturalist; photo examined
EcuadorPastazaPindoyacuOrtega-Andrade 2010
EcuadorPastazaRío Anzu ReserveReyes-Puig et al. 2013
EcuadorPastazaRío BufeoOrtega-Andrade 2010
EcuadorPastazaRío LliquinoZaher 1996
EcuadorPastazaRío PuninoNogueira et al. 2019
EcuadorPastazaRío VillanoNogueira et al. 2019
EcuadorPastazaSan José de CurarayiNaturalist; photo examined
EcuadorPastazaSanta AnaNogueira et al. 2019
EcuadorPastazaSanta ClaraNogueira et al. 2019
EcuadorPastazaVillanoReptiles of Ecuador book database
EcuadorSucumbíosDurenoYánez-Muñoz & Chimbo 2007
EcuadorSucumbíosEl Eno, 4.7 km N ofNogueira et al. 2019
EcuadorSucumbíosEl ReventadorMHNG 2445.005
EcuadorSucumbíosEstación Amazonas OCPValencia & Garzón 2011
EcuadorSucumbíosEstación CayagamaValencia & Garzón 2011
EcuadorSucumbíosEstación PUCE en CuyabenoNogueira et al. 2019
EcuadorSucumbíosGarzacochaiNaturalist; photo examined
EcuadorSucumbíosLa Selva LodgeReptiles of Ecuador book database
EcuadorSucumbíosLagartocochaiNaturalist; photo examined
EcuadorSucumbíosLaguna GrandeNogueira et al. 2019
EcuadorSucumbíosLimoncochaLACM 73327; VertNet
EcuadorSucumbíosMushullactaNogueira et al. 2019
EcuadorSucumbíosPisoriéYánez-Muñoz & Chimbo 2007
EcuadorSucumbíosPozo Lago NorteiNaturalist; photo examined
EcuadorSucumbíosRío MaloNogueira et al. 2019
EcuadorSucumbíosSan Pablo de KantesiyaNogueira et al. 2019
EcuadorSucumbíosSani LodgeReptiles of Ecuador book database
EcuadorSucumbíosSanta CeciliaNogueira et al. 2019
EcuadorSucumbíosTarapoaNogueira et al. 2019
EcuadorTungurahuaCerro Candelaria ReserveReptiles of Ecuador book database
EcuadorTungurahuaReserva Río ZuñacYánez-Muñoz et al. 2013
EcuadorZamora ChinchipeAlto MachinazaNogueira et al. 2019
EcuadorZamora ChinchipeCerro PlateadoDarwin Núñez, pers. comm.
EcuadorZamora ChinchipeCiudad PerdidaOnline multimedia
EcuadorZamora ChinchipeEl Chorro, 1.4 km S ofReptiles of Ecuador book database
EcuadorZamora ChinchipeJamboéDarwin Núñez, pers. comm.
EcuadorZamora ChinchipeJambué BajoNogueira et al. 2019
EcuadorZamora ChinchipeLoja, 21 km E ofZaher 1996
EcuadorZamora ChinchipeReserva Biológica Cerro PlateadoNogueira et al. 2019
EcuadorZamora ChinchipeVia RomerillosDarwin Núñez, pers. comm.
EcuadorZamora ChinchipeZamoraCisneros-Heredia et al. 2007
EcuadorZamora ChinchipeZumba–Pucubamba roadThis work; Fig. 1
PeruAmazonasChiriaco, 43 km NE ofLSUMZ 39325; VertNet
PeruAmazonasCumbaKoch et al. 2018
PeruAmazonasQuebrada HondaKoch et al. 2018
PeruCajamarcaPericoKoch et al. 2018
PeruLoretoParinariNogueira et al. 2019