Published November 29, 2020. Updated April 24, 2024. Open access. | Gallery ❯ |
Golden-bellied Marsh-Snake (Erythrolamprus albiventris)
Reptiles of Ecuador | Serpentes | Colubridae | Erythrolamprus albiventris
English common name: Golden-bellied Marsh-Snake.
Spanish common names: Culebra ventridorada, culebrilla ventridorada, culebra boba de vientre pálido, culebra boba verde.
Recognition: ♂♂ 67.9 cmMaximum distance from the snout to the tip of the tail. ♀♀ 80.5 cmMaximum distance from the snout to the tip of the tail.. Erythrolamprus albiventris is a medium-sized snake having a white or yellow immaculate belly. The dorsum is bright leaf-green or olive with thin black stripes along the posterior half of the body and tail (Fig. 1).1 This species differs from other greenish diurnal snakes in western Ecuador (particularly Chironius exoletus, C. monticola, and Dendrophidion graciliverpa) by being smaller (total length <1 meter), having contrasting black stripes on the body (not only on the tail), and by having smooth dorsal scales arranged in 17 rows at mid-body.2 This species differs from E. fraseri by having immaculate, rather than black-checkered, ventral surfaces.1 Young individuals of E. albiventris usually have a black nape band.
Figure 1: Individuals of Erythrolamprus albiventris from Ecuador: Guanazán, El Oro province (); Santa Lucía Reserve, Pichincha province (); Valle de los Chillos, Pichincha province (); Guápulo, Pichincha province (); Milpe, Pichincha province (); Manduriacu Reserve, Imbabura province (); Otonga Reserve, Cotopaxi province (). ad=adult, j=juvenile.
Natural history: Erythrolamprus albiventris is a comparatively common diurnal and terrestrial snake that inhabits evergreen lowland forests, cloud forests, and humid montane shrublands. The species also occurs in areas having a matrix of pastures, plantations, and remnants of native vegetation, as well as in gardens of heavily-populated urban areas such as Quito.2 Golden-bellied Marsh-Snakes occur in higher densities along marshes, swamps, artificial ponds, and along streams,2,3 and they also venture inside caves and abandoned mines.4 Most active individuals are seen during sunny hours in the morning, crossing roads and trails, basking in open areas or foraging on leaf-litter, soil, or among grass or shrubs.2,3,5 However, individuals are capable of climbing on vegetation up to 2 m above the ground.3 When not active, they remain hidden under logs, timber, agave plants, dirt clods, stones, building blocks, or in crevices.2,3,5
Golden-bellied Marsh-Snakes have an aglyphous dentition,1 meaning their teeth lack specialized grooves to deliver venom. They are active hunters having a diet primarily based on frogs (including Hyloxalus infraguttatus,4 Epipedobates anthonyi,4 Leptodactylus melanonotus,2 Pristimantis achatinus,5 and P. unistrigatus),3 and tadpoles of Gastrotheca riobambae.3 They also feed on lizards such as Gonatodes caudiscutatus,4 Lepidoblepharis conolepis,2 Pholidobolus montium6 and Stenocercus guentheri.7 Individuals are usually calm and try to flee when threatened, relying mostly on crypsis as a primary defense mechanism. If disturbed, they may flatten their body dorsoventrally and produce a musky odor.3 Individuals of Erythrolamprus albiventris are preyed upon by snakes (Erythrolamprus mimus) and by domestic animals (cats and chickens).2
Breeding congregations of Erythrolamprus albiventris have been observed during the rainy season in western Ecuador (December–May). In one hole, five males and one female were coiled into a reproductive ball.5,8 Clutches of 5–10 eggs have been found in heaps of rotten vegetation.9,10
Conservation: Near Threatened Not currently at risk of extinction, but requires some level of management to maintain healthy populations..11 Erythrolamprus albiventris is included in this category on the basis of the species’ wide distribution (here estimated to be ~53,539 km2), presence in over a dozen protected areas, and presumed stable populations.2 The main threat to the long-term survival of populations of E. albiventris is the continuing decline in the extent and quality of its habitat, mostly due to encroaching human activities such as agriculture, cattle grazing, wild fires, and the replacement of native vegetation with eucalyptus and pine trees. Approximately 63.3% of the potential habitat of the species in Ecuador has already been destroyed.12 Golden-bellied Marsh-Snakes also suffer from human persecution and traffic-related mortality.3,5 Therefore, the species may qualify for a threatened category in the near future if these threats are not addressed. However, there is no current information on the population trend of E. albiventris to determine whether its numbers are declining.
Distribution: Erythrolamprus albiventris is endemic to an area of approximately 53,539 km2 in Ecuador, where it is distributed throughout the Chocoan lowlands and adjacent foothills of the Andes as well as in the inter-Andean valley of Quito (Fig. 2).
Figure 2: Distribution of Erythrolamprus albiventris in Ecuador. See Appendix 1 for a complete list of the presence localities included in the map.
Etymology: The name Erythrolamprus, which comes from the Greek words erythros (=red) and lampros (=brilliant),13 refers to the bright red body rings of some snakes in this genus (such as E. aesculapii). The specific epithet albiventris, which comes from the Latin words albus (=white) and venter (=belly),13 refers to the immaculate ventral surfaces.14
See it in the wild: Individuals of Erythrolamprus albiventris are frequently encountered in forested areas throughout the specie’s distribution. However, they are particularly abundant in the general area of Mindo and along the old Nono–Mindo road, Pichincha province. The snakes may be spotted as they cross trails and roads in areas having adequate vegetation cover, especially during sunny mornings.
Acknowledgments: Special thanks to Carlos Zorrilla, Eduardo Zavala, and Jose Manuel Falcón for providing locality data and natural history information for Erythrolamprus albiventris. This account was published with the support of Secretaría Nacional de Educación Superior Ciencia y Tecnología (programa INEDITA; project: Respuestas a la crisis de biodiversidad: la descripción de especies como herramienta de conservación; No 00110378), Programa de las Naciones Unidas (PNUD), and Universidad San Francisco de Quito (USFQ).
Special thanks to Annalaura Averill-Murray for symbolically adopting the Golden-bellied Marsh-Snake and helping bring the Reptiles of Ecuador book project to life.
Click here to adopt a species.
Author: Alejandro ArteagaaAffiliation: Khamai Foundation, Quito, Ecuador.
Photographer: Jose VieiraaAffiliation: Tropical Herping (TH), Quito, Ecuador.,bAffiliation: ExSitu, Quito, Ecuador.
How to cite? Arteaga A (2024) Golden-bellied Marsh-Snake (Erythrolamprus albiventris). 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/LLLF9348
Literature cited:
- Dixon JR (1983) Systematics of the Latin American snake Liophis epinephelus (Serpentes: Colubridae). In: Rhodin AGJ, Miyamata K (Eds) Advances in herpetology and evolutionary biology. Museum of Comparative Zoology, Cambridge, 132–149.
- Arteaga A, Bustamante L, Guayasamin JM (2013) The amphibians and reptiles of Mindo. Universidad Tecnológica Indoamérica, Quito, 257 pp.
- Ramírez-Jaramillo S (2015) Observaciones sobre la historia natural de Erythrolamprus epinephelus albiventris en el valle de quito, Ecuador. Avances en Ciencias e Ingenierías 7: 5–7.
- Jose Manuel Falcón, pers. comm.
- Field notes, Reptiles of Ecuador book project.
- Mafla-Endara P, Ayala-Varela F (2012) Pholidobolus montium (lagartija minadora). Predation. Herpetological Review 43: 137.
- Cadena-Ortiz H, Barahona A, Bahamonde-Vinueza D, Brito J (2017) Anecdotal predation events of some snakes in Ecuador. Herpetozoa 30: 93–96.
- Photographic record by Silvio Paladines.
- Morales MA (2004) Dinámica poblacional de las comunidades de anfibios y reptiles de siete localidades de la zona de amortiguamiento de la Reserva Ecológica Cotacachi-Cayapas, Esmeraldas, Ecuador. Universidad del Azuay, 122 pp.
- Copping R (1957) Reptiles and amphibians of the highlands of Ecuador. British Journal of Herpetology 2: 45–56.
- Carrillo E, Aldás A, Altamirano M, Ayala F, Cisneros-Heredia DF, Endara A, Márquez C, Morales M, Nogales F, Salvador P, Torres ML, Valencia J, Villamarín F, Yánez-Muñoz M, Zárate P (2005) Lista roja de los reptiles del Ecuador. Fundación Novum Millenium, Quito, 46 pp.
- MAE (2012) Línea base de deforestación del Ecuador continental. Ministerio del Ambiente del Ecuador, Quito, 30 pp.
- Brown RW (1956) Composition of scientific words. Smithsonian Books, Washington D.C., 882 pp.
- Jan G (1863) Enumerazione sistematica degli ofidi appartenenti al gruppo Coronellidae. Archive per la Zoologia, l’Anatomia et la Fisiologia 2: 215–330.
Appendix 1: Locality data used to create the distribution map of Erythrolamprus albiventris in Ecuador (Fig. 2). Go to the section on symbols and abbreviations for a list of acronyms used.
| Country | Province | Locality | Source |
| Ecuador | Azuay | Chilcaplaya | José Manuel Falcón, pers. comm. |
| Ecuador | Azuay | Los Monos | Reptiles of Ecuador book database |
| Ecuador | Azuay | Sarayunga | José Manuel Falcón, pers. comm. |
| Ecuador | Azuay | Vall de Chaucha | Dixon 1983 |
| Ecuador | Azuay | Yunguilla | José Manuel Falcón, pers. comm. |
| Ecuador | Bolívar | Balzapamba | Dixon 1983 |
| Ecuador | Bolívar | Chazo Juan | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Bolívar | Cumandá, 7 km NW of | iNaturalist; photo examined |
| Ecuador | Cañar | Finca Huatacón | Reptiles of Ecuador book database |
| Ecuador | Cañar | La Troncal | MZUA.RE.0038; examined |
| Ecuador | Carchi | Chical | USNM 286302; VertNet |
| Ecuador | Chimborazo | Valle del Chanchán | ANSP 18119; VertNet |
| Ecuador | Cotopaxi | Bosque Integral Otonga | This work; Fig. 1 |
| Ecuador | Cotopaxi | Cutzualo | MHNG 2442.053; collection database |
| Ecuador | Cotopaxi | Galápagos | MHNG 2411.041; collection database |
| Ecuador | Cotopaxi | Las Damas | MHNG 2458.021; collection database |
| Ecuador | Cotopaxi | Las Pampas | César Tapia, pers. comm. |
| Ecuador | Cotopaxi | Naranjito | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Cotopaxi | Palo Quemado | MHNG 2410.100; collection database |
| Ecuador | Cotopaxi | Peñas Coloradas, Sigchos | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | El Oro | Bella María | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | El Oro | Chilla | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | El Oro | Chivaturco | Garzón-Santomaro et al. 2019 |
| Ecuador | El Oro | El Progreso | MHNG 2308.078; collection database |
| Ecuador | El Oro | Guanazán | This work; Fig. 1 |
| Ecuador | El Oro | Machala, 10 km SE of | Dixon 1983 |
| Ecuador | El Oro | Reserva Biológica Buenaventura | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | El Oro | Urna de Buenaventura, 4 km W | iNaturalist; photo examined |
| Ecuador | El Oro | Zambotambo | Reptiles of Ecuador book database |
| Ecuador | Esmeraldas | Atacames | MHNG 2221.057; collection database |
| Ecuador | Esmeraldas | Centro de Fauna Silvestre James Brown | Photo by Salvador Palacios |
| Ecuador | Esmeraldas | Cube, 1.7 km E of | iNaturalist; photo examined |
| Ecuador | Esmeraldas | Esmeraldas | USNM 232824; VertNet |
| Ecuador | Esmeraldas | Mayronga, Lagarto | Torres-Carvajal et al. 2019 |
| Ecuador | Esmeraldas | Pajonal | Morales 2004 |
| Ecuador | Esmeraldas | Playa Rica | MNHN 1902.356; collection database |
| Ecuador | Esmeraldas | Reserva Biológica Bilsa | Ortega-Andrade et al. 2010 |
| Ecuador | Esmeraldas | Reserva Canandé | Yánez-Muñoz et al. 2005 |
| Ecuador | Esmeraldas | Viche | Hinojosa-Almeida 2021 |
| Ecuador | Guayas | Balzar | Dixon 1983 |
| Ecuador | Guayas | Bucay | Photo by Keyko Cruz |
| Ecuador | Guayas | Capeira | Photo by Eduardo Zavala |
| Ecuador | Guayas | Taura | iNaturalist; photo examined |
| Ecuador | Imbabura | Cabañas Colibrí | iNaturalist; photo examined |
| Ecuador | Imbabura | El Refugio de Intag | Photo by Peter Joost |
| Ecuador | Imbabura | El Rosario, 2 km E of | iNaturalist; photo examined |
| Ecuador | Imbabura | Intag | Peter Joost, pers. comm. |
| Ecuador | Imbabura | Intag Cloudforest Reserve | Photo by Carlos Zorrilla |
| Ecuador | Imbabura | La Merced de Buenos Aires | Jesse Kimmerling, pers. comm. |
| Ecuador | Imbabura | Lita | USNM 232905; VertNet |
| Ecuador | Imbabura | Paramba | Boulenger 1898 |
| Ecuador | Imbabura | Reserva Los Cedros | Photo by Fernando Rojas |
| Ecuador | Imbabura | Reserva Manduriacu | This work; Fig. 1 |
| Ecuador | Imbabura | Reserva Siempre Verde | iNaturalist; photo examined |
| Ecuador | Imbabura | Río Aguas Claras | Dixon 1983 |
| Ecuador | Loja | Alamor | Dixon 1983 |
| Ecuador | Loja | Alamor, vicinity of | Reptiles of Ecuador book database |
| Ecuador | Loja | Camino a Vicentino | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Loja | Celica | Photo by Jorge Castillo |
| Ecuador | Loja | El Limo | iNaturalist; photo examined |
| Ecuador | Loja | Huajala | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Loja | San Ramón | USNM 232906; VertNet |
| Ecuador | Loja | Y de Balsones | iNaturalist; photo examined |
| Ecuador | Los Ríos | Centro Científico Río Palenque | KU 152605; VertNet |
| Ecuador | Los Ríos | Hacienda La Clementina | Dixon 1983 |
| Ecuador | Los Ríos | Río Baba | AMNH 110588; VertNet |
| Ecuador | Los Ríos | Río Baba, 24 km S of | Dixon 1983 |
| Ecuador | Manabí | Vía a San Isidro | Photo by Regdy Vera |
| Ecuador | Pichincha | Alambi | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Aldea Ecoturística Puma Sacha | iNaturalist; photo examined |
| Ecuador | Pichincha | Alluriquín | Dixon 1983 |
| Ecuador | Pichincha | Amaguaña | Hinojosa-Almeida 2021 |
| Ecuador | Pichincha | Barrio Runahurco | iNaturalist; photo examined |
| Ecuador | Pichincha | Bellavista Cloudforest Reserve | Pascal Vagner |
| Ecuador | Pichincha | Below Pacto | Dixon 1983 |
| Ecuador | Pichincha | Birdwatcher House | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Bosque Protector Verdecocha | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Bosque Seco Nueva Esperanza–Guayllabamba | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Calacali–Nanegalito road | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Carcelén | Valencia et al. 2017 |
| Ecuador | Pichincha | Carcelén Bajo | Valencia et al. 2017 |
| Ecuador | Pichincha | Carretas | Valencia et al. 2017 |
| Ecuador | Pichincha | Cayambe | MHNG 2399.099; collection database |
| Ecuador | Pichincha | Cerro Montecristi | Valencia et al. 2017 |
| Ecuador | Pichincha | Checa | iNaturalist; photo examined |
| Ecuador | Pichincha | Chiriboga | MHNG 2221.055; collection database |
| Ecuador | Pichincha | Club Los Chillos | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Pichincha | Dos Puentes | Photo by René Lima |
| Ecuador | Pichincha | Dos Ríos, 3.7 km NE of | KU 142807; VertNet |
| Ecuador | Pichincha | Downtown Mindo | Reptiles of Ecuador book database |
| Ecuador | Pichincha | El Cedral | Reptiles of Ecuador book database |
| Ecuador | Pichincha | El Pancillo | Valencia et al. 2017 |
| Ecuador | Pichincha | El Quinche | Arteaga et al. 2013 |
| Ecuador | Pichincha | El Tingo | MHNG 2442.054; collection database |
| Ecuador | Pichincha | Estación La Favorita | iNaturalist; photo examined |
| Ecuador | Pichincha | Gualea | Dixon 1983 |
| Ecuador | Pichincha | Guápulo | This work; Fig. 1 |
| Ecuador | Pichincha | Hacienda Las Palmas-Río Blanco | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Hacienda Olalla | Dixon 1983 |
| Ecuador | Pichincha | Ilaló | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Inga | Arteaga et al. 2013 |
| Ecuador | Pichincha | Kapari | Reptiles of Ecuador book database |
| Ecuador | Pichincha | La Armenia | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Pichincha | La Hesperia | Brouwer 2018 |
| Ecuador | Pichincha | La Unión–Río Cinto | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Las Palmas | MCZ 164433; VertNet |
| Ecuador | Pichincha | Llambo | Dixon 1983 |
| Ecuador | Pichincha | Los Bancos | iNaturalist; photo examined |
| Ecuador | Pichincha | Machachi | Arteaga et al. 2013 |
| Ecuador | Pichincha | Malchinguí | Photo by José Campaña |
| Ecuador | Pichincha | Mashpi | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Milpe | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Mindo Lindo | Photo by Heike Brieschke |
| Ecuador | Pichincha | Mindo–Cunuco | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Mindo–Las Cascadas | Photo by David Hamill |
| Ecuador | Pichincha | Mindo, 4 km N of | Dixon 1983 |
| Ecuador | Pichincha | Molinuco | iNaturalist; photo examined |
| Ecuador | Pichincha | Nanegalito | MHNG 2442.057 |
| Ecuador | Pichincha | Nanegalito–Mindo | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Nanegalito, 1.5 km E of | Dixon 1983 |
| Ecuador | Pichincha | Near Nanegalito | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Nono | Photo by Jaime Cháves |
| Ecuador | Pichincha | Nono–Mindo road | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Nono, 13 km NW of | KU 158534; VertNet |
| Ecuador | Pichincha | Nono, 4.5 km NW of | KU 218423; VertNet |
| Ecuador | Pichincha | Nuevo Aeropuerto | Photo by Eduardo Toral |
| Ecuador | Pichincha | Pachijal | Photo by Vinicio Perez |
| Ecuador | Pichincha | Parque de la Mujer | Valencia et al. 2017 |
| Ecuador | Pichincha | Parque Itchimbía | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Parque Los Mastodontes | Valencia et al. 2017 |
| Ecuador | Pichincha | Parque Metropolitano | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Pifo | Arteaga et al. 2013 |
| Ecuador | Pichincha | Pintag | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Puerto Quito | MCZ 164502; VertNet |
| Ecuador | Pichincha | Pulhulahua Reserve | iNaturalist; photo examined |
| Ecuador | Pichincha | Quito, 12 de Octubre | Torres-Carvajal & Hinojosa 2020 |
| Ecuador | Pichincha | Quito, 15 km NE of | Dixon 1983 |
| Ecuador | Pichincha | Quito, Colegio San Gabriel | Valencia et al. 2017 |
| Ecuador | Pichincha | Quito, Jardines del Batán | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Reserva Las Gralarias | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Reservorio Cumbayá | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Río Blanco | Dixon 1983 |
| Ecuador | Pichincha | Río Cinto | Photo by Lisa Brunetti |
| Ecuador | Pichincha | Río Mindo | Dixon 1983 |
| Ecuador | Pichincha | Río Orito | Torres-Carvajal & Salazar-Valenzuela 2012 |
| Ecuador | Pichincha | Río Saloya | Dixon 1983 |
| Ecuador | Pichincha | Río Silanche | Dixon 1983 |
| Ecuador | Pichincha | San Rafael | Ramírez-Jaramillo 2015 |
| Ecuador | Pichincha | San Roque | Valencia et al. 2017 |
| Ecuador | Pichincha | Santa Lucía Alta | iNaturalist; photo examined |
| Ecuador | Pichincha | Santa Lucía Reserve | Savit 2006 |
| Ecuador | Pichincha | Saragoza–Río Cinto | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Séptimo Paraíso | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Tambo Tanda | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Tamboquinde | Yánez-Muñoz et al. 2009 |
| Ecuador | Pichincha | Tandapi | KU 112272; VertNet |
| Ecuador | Pichincha | Tandapi, 9 km SE of | AMNH 113020; VertNet |
| Ecuador | Pichincha | Tandayapa | MHNG 2444.055; collection database |
| Ecuador | Pichincha | Tanti | Boulenger 1884 |
| Ecuador | Pichincha | Terrabambú | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Terreno de Eric Osterman en Mindo | Reptiles of Ecuador book database |
| Ecuador | Pichincha | Tola Chica | Photo by Morley Read |
| Ecuador | Pichincha | Tumbaco | Boulenger 1882 |
| Ecuador | Pichincha | Valle de los Chillos | This work; Fig. 1 |
| Ecuador | Pichincha | Way to Mashpi | iNaturalist; photo examined |
| Ecuador | Pichincha | Yaruquí | Arteaga et al. 2013 |
| Ecuador | Pichincha | Yellow House | Reptiles of Ecuador book database |
| Ecuador | Santo Domingo de los Tsáchilas | Bosque Protector La Perla | Photo by Plácido Palacios |
| Ecuador | Santo Domingo de los Tsáchilas | Bosque Protector Río Guajalito | Reptiles of Ecuador book database |
| Ecuador | Santo Domingo de los Tsáchilas | Chiriboga, 4 km W of | KU 142806; VertNet |
| Ecuador | Santo Domingo de los Tsáchilas | Chiriboga, 8 km W of | Dixon 1983 |
| Ecuador | Santo Domingo de los Tsáchilas | La Favorita | Reyes-Puig et al. 2015 |
| Ecuador | Santo Domingo de los Tsáchilas | Las Palmeras | AMNH 6042; VertNet |
| Ecuador | Santo Domingo de los Tsáchilas | Miligali | Boulenger 1883 |
| Ecuador | Santo Domingo de los Tsáchilas | Río Mulaule | Dixon 1983 |
| Ecuador | Santo Domingo de los Tsáchilas | Santo Domingo | Dixon 1983 |
| Ecuador | Santo Domingo de los Tsáchilas | Santo Domingo, 5 km SSW of | Dixon 1983 |