Research Article |
Corresponding author: Omar Torres-Carvajal ( omartorcar@gmail.com ) Academic editor: Johannes Penner
© 2015 Omar Torres-Carvajal, Pablo J. Venegas, Kevin de Queiroz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Torres-Carvajal O, Venegas PJ, de Queiroz K (2015) Three new species of woodlizards (Hoplocercinae, Enyalioides) from northwestern South America. ZooKeys 494: 107-132. https://doi.org/10.3897/zookeys.494.8903
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The discovery of three new species of Enyalioides from the tropical Andes in Ecuador and northern Peru is reported. Enyalioides altotambo sp. n. occurs in northwestern Ecuador and differs from other species of Enyalioides in having dorsal scales that are both smooth and homogeneous in size, a brown iris, and in lacking enlarged, circular and keeled scales on the flanks. Enyalioides anisolepis sp. n. occurs on the Amazonian slopes of the Andes in southern Ecuador and northern Peru and can be distinguished from other species of Enyalioides by its scattered, projecting large scales on the dorsum, flanks, and hind limbs, as well as a well-developed vertebral crest, with the vertebrals on the neck at least three times higher than those between the hind limbs. Enyalioides sophiarothschildae sp. n. is from the Amazonian slopes of the Cordillera Central in northeastern Peru; it differs from other species of Enyalioides in having caudal scales that are relatively homogeneous in size on each caudal segment, a white gular region with a black medial patch and several turquoise scales in males, as well as immaculate white labials and chin. A molecular phylogenetic tree of 18 species of hoplocercines is presented, including the three species described in this paper and E. cofanorum, as well as an updated identification key for species of Hoplocercinae.
Reportamos el descubrimiento de tres especies nuevas de Enyalioides de los Andes tropicales en Ecuador y norte de Perú. Enyalioides altotambo sp. n., del noroccidente de Ecuador, difiere de otras especies de Enyalioides por poseer escamas dorsales lisas y homogéneas en tamaño, iris café y por carecer de escamas circulares grandes y quilladas en los flancos. Enyalioides anisolepis sp. n. ocurre en las estribaciones amazónicas de los Andes al sur de Ecuador y norte de Perú, y se distingue de otras especies de Enyalioides por poseer escamas grandes y proyectadas dispersas en el dorso, flancos y extremidades posteriores, así como por su cresta vertebral bastante desarrollada, que a nivel del cuello es tres veces más alta que entre las extremidades posteriores. Enyalioides sophiarothschildae sp. n., de las estribaciones amazónicas de la Cordillera Central al norte del Perú, difiere de otras especies de Enyalioides por poseer escamas caudales de tamaño similar en cada segmento caudal, una región gular blanca con una mancha medial negra y escamas turquesa en machos, así como la quijada y labiales de color blanco. También presentamos un árbol filogenético molecular de 18 especies de hoplocercinos, que incluye a las tres especies descritas en este artículo y a E. cofanorum, así como una clave de identificación actualizada para las especies de Hoplocercinae.
Andes, Ecuador, Enyalioides, Hoplocercinae, Iguania, lizards, new species, Peru, systematics
The iguanian lizard clade Hoplocercinae includes 16 currently recognized species assigned to Enyalioides, Hoplocercus, and Morunasaurus distributed from Panama to central Brazil (
With nearly 40% of the total number of species described in the last seven years from Ecuador and Peru (
Snout-vent length (SVL) and tail length (TL) measurements were made with a ruler and recorded to the nearest millimeter. All other measurements (i.e., head width, length and height; rostral and mental width and height) were made with digital calipers and recorded to the nearest 0.1 mm. Sex was determined by noting the presence of hemipenes or sexually dichromatic characters. The format of
Following laboratory protocols similar to those presented by
Vouchers, locality data, and GenBank accession numbers of new DNA sequences obtained for this study.
Taxon | Voucher | Locality | GenBank number (ND4) | GenSeq nomenclature |
---|---|---|---|---|
E. altotambo | QCAZ 8073 (holotype) | Ecuador: Esmeraldas: Alto Tambo, 5 km on road to Placer | KP235211 | genseq-1 |
E. anisolepis | QCAZ 8395 | Ecuador: Zamora-Chinchipe: Chito, sector Los Planes | KP235213 | genseq-2 |
E. anisolepis | QCAZ 8428 | Ecuador: Zamora-Chinchipe: Chito | KP235214 | genseq-2 |
E. anisolepis | QCAZ 8515 | Ecuador: Zamora-Chinchipe: Chito, sector Los Planes | KP235215 | genseq-2 |
E. cofanorum | QCAZ 8035 | Ecuador: Orellana: 66 km on road Pompeya-Iro | KP235210 | genseq-4 |
E. sophiarothschildae | CORBIDI 647 (holotype) | Peru: San Martín: Río Lejía on the trail La Cueva-Añazco Pueblo | KP235212 | genseq-1 |
Editing, assembly, and alignment of sequences were performed with Geneious 7.1.7 (
Phylogenetic relationships were assessed under a Bayesian inference approach using MrBayes 3.2.2 (
The taxonomic conclusions of this study are based on the observation of morphological features and color patterns, as well as the inferred phylogenetic relationships. This information is considered as species delimitation criteria following a general lineage or unified species concept (
Enyalioides oshaughnessyi (part)
Holotype. QCAZ 8073 (Fig.
Paratype. ECUADOR: Provincia Esmeraldas: QCAZ 6671, adult female, Alto Tambo, Balthazar river, 0.90000°N; -78.61667°W, 645 m, collected on 5 November 2005 by F. Ayala-Varela and I.G. Tapia.
Enyalioides altotambo differs from other species of Enyalioides, except for E. oshaughnessyi, in having dorsal scales that are both smooth and homogeneous in size. It can be distinguished from E. oshaughnessyi (character states in parentheses) by the following characters: iris brown in both sexes (iris bright red in both sexes); scales on lateral edge of skull roof just posterior to superciliaries strongly projected (moderately projected); adults of both sexes with light green spots on dorsum (if present, spots turquoise or blue); adult males with a black medial patch on gular region not extending dorsally to form an antehumeral bar (black patch under gular fold extending dorsally to form a short antehumeral bar); scales on flanks almost homogenous in size (flank scales heterogeneous in size, with a few enlarged, circular, keeled scales); pale postympanic stripe on lateral aspect of neck in both sexes (pale postympanic spot in both sexes), posterior surface of thighs without enlarged scales (scattered enlarged scales), tail length/total length 0.57–0.60 (0.59–0.62).
Male (Fig.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least four times higher than those between hind limbs; crest bifurcates at a point approximately 10 mm posterior to the cloaca, and extends onto tail about 1/3 its length; body flanks between fore and hind limbs with slight dorsolateral fold; scales on dorsolateral fold slightly larger than adjacent scales; dorsal and flank scales small, smooth, imbricate, more or less homogeneous in size; ventral scales imbricate, keeled, rectangular or rhomboid, with a posterolateral mucron; ventrals more than twice the size (area) of dorsals.
Limb scales keeled dorsally and keeled or feebly keeled ventrally; scales on dorsal and posterior aspects of thighs heterogeneous in size, with most scales less than half the size of those scales on anterior and ventral aspects, separated from each other by skin covered with tiny granular scales; subdigitals on finger IV 25; subdigitals on toe IV 29; femoral pores on each side one; tail laterally compressed and gradually tapering posteriorly; caudal scales smooth at the base of tail, becoming keeled and imbricate towards tip, gradually increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and seven scales long dorsally.
(Fig.
Variation in meristic and morphometric characters of Enyalioides altotambo are presented in Table
Summary of morphological characters and measurements (mm) of Enyalioides altotambo, E. anisolepis and E. sophiarothschildae. Range (first line) and mean ± standard deviation (second line) are given for quantitative characters, except when there was no variation.
Character | E. altotambo N = 2 | E. anisolepis N = 15 | E. sophiarothschildae N = 3 |
---|---|---|---|
Dorsals in transverse row between dorsolateral crests at midbody | 39–40 39.5 ± 0.71 |
28–35 32.00 ± 2.83 |
22–26 24.33 ± 2.08 |
Ventrals in transverse row at midbody | 31–33 32.00 ± 1.41 |
23–29 26.53 ± 1.92 |
23–26 25.00 ± 1.73 |
Vertebrals from occiput to base of tail | 50–51 50.50 ± 0.71 |
43–62 50.87 ± 6.27 |
51–57 54.00 ± 3.00 |
Gulars | 47 | 30–35 31.71 ± 1.49 |
36 |
Infralabials | 11 | 9 | 9–11 10.00 ± 1.00 |
Supralabials | 13 | 10–12 10.77 ± 0.60 |
9–12 10.67 ± 1.53 |
Canthals | 5 | 5–6 5.43 ± 0.51 |
5 |
Superciliaries | 14–17 15.50 ± 2.12 |
13–18 15.57 ± 1.40 |
13–15 14.00 ± 1.00 |
Transverse rows of ventrals between fore and hind limbs | 47–49 48.00 ± 1.41 |
38–46 41.27 ± 2.60 |
37–40 38.67 ± 1.53 |
Subdigitals finger IV | 23–25 24.00 ± 1.41 |
15–20 18.36 ± 1.39 |
18–19 18.67 ± 0.58 |
Subdigitals toe IV | 27–29 28.00 ± 1.41 |
24–27 25.14 ± 0.86 |
22–27 25.33 ± 2.89 |
Femoral pores | 1–2 1.50 ± 0.71 |
0–3 1.64 ± 1.01 |
3–4 3.67 ± 0.58 |
Tail length/Total length | 0.57–0.60 0.59 ± 0.02 |
0.59–0.71 0.62 ± 0.03 |
0.60–0.61 0.61 ± 0.01 |
Enyalioides altotambo is only known from two adjacent localities at 620–645 m in the Chocoan rainforests of northwestern Ecuador (Fig.
The specific epithet is a noun in apposition and refers to Alto Tambo, Provincia Esmeraldas, Ecuador, a village on the Ibarra-San Lorenzo road where Enyalioides altotambo was discovered.
Although previously referred to Enyalioides oshaughnessyi, the possibility that the specimens here named Enyalioides altotambo represented a distinct species was recognized in previous studies. In a phylogenetic analysis of hoplocercine lizards,
Holotype. QCAZ 12537 (Fig.
Paratypes (14). ECUADOR: Provincia Zamora-Chinchipe: QCAZ 12521, juvenile with the same collection data as the holotype except -4.88673°S, -79.08744°W, 738 m; QCAZ 12527, adult male (Fig.
Enyalioides anisolepis can be distinguished from other species of Enyalioides, except for E. heterolepis, by having conical dorsal head scales (only in E. anisolepis and E. heterolepis) and scattered, projecting, large scales on the dorsum, flanks, and hind limbs (also in E./Morunasaurus annularis and E./M. groi), which are conspicuous in adults of both sexes (Fig.
The only other species of Enyalioides with scattered, projecting dorsal scales is E. cofanorum, which differs from E. anisolepis in lacking projecting scales on the hind limbs, and in being smaller in size (maximum SVL in males and females of E. cofanorum 107 mm and 109 mm, respectively; 130 mm and 119 mm in E. anisolepis). Additionally, adults of both sexes of E. cofanorum have a brownish background (marked sexual dichromatism in E. anisolepis).
Male (Fig.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least three times higher than those between hind limbs; crest bifurcates at a point approximately 10 mm posterior to the cloaca, and extends onto tail about ¼ its length; flanks between fore and hind limbs with dorsolateral and ventrolateral longitudinal folds, as well as several oblique folds; axillary region with three vertical folds; scales on dorsolateral folds slightly larger than adjacent scales giving the fold the appearance of a low crest; scales between dorsolateral folds and vertebral crest heterogeneous in size, prominently keeled, and imbricate, with largest scales twice as large as smallest ones; neck and scapular region with scattered, large conical scales; flank scales ventral to dorsolateral folds similar to those dorsal to folds, with largest scales four times as large as smallest ones (Fig.
Limb scales keeled dorsally and ventrally, homogeneous in size on fore limbs; scales on dorsal and posterior aspect of thighs heterogeneous in size, with most scales less than half the size of those on anterior and ventral aspects; scales on dorsal surface of shanks heterogeneous in size, with granular scales between large keeled scales; subdigitals on finger IV 17; subdigitals on toe IV 25; three femoral pores on left leg, two on right leg; tail laterally compressed and gradually tapering posteriorly; caudal scales strongly keeled and imbricate, increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and six scales long dorsally.
(Fig.
Variation in meristic and morphometric characters of Enyalioides anisolepis are presented in Table
This species has a marked sexual dichromatism in background colors (green in males, brown in females; see Fig.
Adult females share similar color patterns with juveniles (Fig.
Enyalioides anisolepis is known to occur between 724–1742 m on the Amazonian slopes of the Andes in southern Ecuador and northern Peru (Fig.
The specific epithet anisolepis is a noun (in apposition) in the nominative singular and derives from the Greek words anisos (= unequal) and lepis (= scale). It refers to the heterogeneous scales on the dorsum, flanks and hind limbs of lizards of this species.
Holotype. CORBIDI 647 (Fig.
Paratypes (2). PERU: Región San Martín: Provincia Mariscal Cáceres: MUSM 21883-84, adult males, El Dorado, -6.76666°S; -77.54500°W, 1600m, collected on 5 December 2003 by P.J. Venegas.
Enyalioides sophiarothschildae can be distinguished from other species of Enyalioides, except for E. laticeps, by having caudal scales that are relatively homogeneous in size on each caudal segment; in all other species of Enyalioides, the dorsal and lateral caudals increase in size posteriorly on each caudal segment, and the largest (posteriormost) caudals on each segment are mucronate or have some kind of projection (
Male (Fig.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least four times higher than those between hind limbs; crest bifurcates posteriorly and extends onto tail less than ¼ its length; body flanks between fore and hind limbs with slight dorsolateral and ventrolateral longitudinal folds; scales on dorsolateral folds similar in size to adjacent scales; dorsal and flank scales small, keeled, imbricate, more or less homogeneous in size, and separated from each other by skin covered with tiny granular scales; ventral scales imbricate, smooth or slightly keeled, rectangular or rhomboid, with a posterolateral mucron; ventrals more than twice the area of dorsals.
Limb scales keeled dorsally and smooth or slightly keeled ventrally; scales on dorsal and posterior surfaces of thighs heterogeneous in size, with most scales less than half the size of those on anterior and ventral surfaces, separated from each other by skin covered with tiny granular scales; subdigitals on finger IV 17; subdigitals on toe IV 25; femoral pores on each side four; tail laterally compressed and gradually tapering posteriorly; caudal scales strongly keeled and imbricate, not gradually increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and six scales long dorsally.
(Fig.
Variation in meristic and morphometric characters of Enyalioides sophiarothschildae are presented in Table
Enyalioides sophiarothschildae is known from the northeastern slopes of the Cordillera Central in Peru between 1600–1700 m (Fig.
Individuals of Enyalioides sophiarothschildae were found active by day in primary forest. The holotype was found crossing a trail and tried to hide between the roots of a big tree when approached for capture. One of the paratypes climbed up a tree three meters above the ground when approached. The other paratype was found sitting on a big root.
The specific epithet is a noun in the genitive case and is a patronym honoring Sophia Rothschild in recognition of her financial support for the improvement of the herpetological collection of CORBIDI through the BIOPAT Program.
The phylogenetic tree inferred in this study (Fig.
50% Majority rule consensus tree of hoplocercine lizards (E. = Enyalioides, M. = Morunasaurus) based on a Bayesian analysis of mtDNA sequences. Posterior probabilities are equal to 1, unless otherwise noted by numbers next to branches. Outgroup taxa are not shown. The notation E./M. indicates that according to the phylogenetic definitions (
Enyalioides anisolepis is strongly supported (PP = 1) as monophyletic and is sister (PP = 0.99) to a clade (PP = 1) composed of E. cofanorum, E. microlepis, E. rubrigularis, and E. praestabilis. Enyalioides sophiarothschildae is sister (PP = 1) to a clade (PP = 1) composed of two recently discovered species, E. binzayedi and E. rudolfarndti (
The following key is artificial in the sense that its structure does not necessarily reflect the order of branching in the phylogeny.
1 | Dorsal head scales flat, smooth, juxtaposed; vertebral crest absent or composed of a discontinuous row of enlarged scales that are longer than tall | 2 |
– | Dorsal head scales conical; vertebral crest present, composed of projecting scales that are taller than long | 5 |
2 | Tail depressed, short (tail length < snout-vent length), with enlarged spiny scales dorsally and laterally | Hoplocercus spinosus |
– | Tail nearly round, moderate (tail length > snout-vent length), with rings of enlarged spiny scales | 3 |
3 | Vertebral region of trunk without enlarged scales; tail with three scale rows separating the spiny whorls ventrally | Morunasaurus groi |
– | Some vertebral scales in trunk region enlarged forming a discontinuous longitudinal row; tail with two scale rows separating the spiny whorls ventrally | 4 |
4 | Usually two femoral pores on each leg; two postmentals; females without streaks on throat | M. annularis |
– | Femoral pores 3–4 on each leg; usually four postmentals; females with dark streaks on throat | M. peruvianus |
5 | Caudal scales homogeneous in size within each autotomic segment | 6 |
– | Caudal scales increase in size posteriorly within each autotomic segment | 7 |
6 | Gular region in males white with a black medial patch | Enyalioides sophiarothschildae |
– | Gular region in males orange or dirty cream, with longitudinal brown, reddish-brown, bluish, or orange streaks, and a large brown or black medial blotch at the level of the gular fold | E. laticeps |
7 | Lateral superciliary projection present; vertebral crest usually discontinuous (absent on posterior part of neck) | E. palpebralis |
– | Lateral superciliary projection absent; vertebral crest continuous | 8 |
8 | Scattered, conspicuous large scales on dorsum, flanks, and hind limbs present | 9 |
– | Scattered, conspicuous large scales on dorsum, flanks, and hind limbs absent | 10 |
9 | Scattered large scales tetrahedral in shape; vertebrals on neck maximum twice as high as those between hind limbs | E. heterolepis |
– | Scattered large scales strongly keeled, not tetrahedral in shape; vertebrals on neck at least three times higher than those between hind limbs | E. anisolepis |
10 | Ventrals smooth or slightly keeled | 11 |
– | Ventrals conspicuously keeled | 12 |
11 | Gulars in males cream or yellow without black margins; usually one femoral pore on each leg | E. praestabilis |
– | Gulars in males bright orange or red, with black margins; usually two femoral pores on each leg | E. rubrigularis |
12 | Dorsals heterogeneous in size, with scattered, tetrahedral, projecting scales (sometimes absent in males or juveniles); dorsolateral crests well developed between hind limbs | E. cofanorum |
– | Dorsals homogeneous in size, without projecting scales; dorsolateral crests inconspicuous or absent between hind limbs | 13 |
13 | Dorsals smooth or slightly keeled | 14 |
– | Dorsals conspicuously keeled | 15 |
14 | Scales on flanks heterogeneous in size, with a few enlarged, circular, keeled scales; iris bright red in both sexes; black patch under gular fold extending dorsally to form a short antehumeral bar in males | E. oshaughnessyi |
– | Scales on flanks almost homogenous in size; iris brown in both sexes; black medial patch on gular region not extending dorsally to form an antehumeral bar in males | E. altotambo |
15 | Dorsals in transverse row between dorsolateral crests at midbody 31 or fewer | 16 |
– | Dorsals in transverse row between dorsolateral crests at midbody more than 31 | 17 |
16 | Scales along the lateral edge of the skull roof strongly projected; dorsal scales homogeneous in size, with prominent median keel; antehumeral orange blotch in adult males absent | E. binzayedi |
– | Scales along the lateral edge of the skull roof slightly projected; dorsal scales heterogeneous in size, without prominent median keel; distinct antehumeral orange blotch in adult males | E. rudolfarndti |
17 | White or cream spot posterior to tympanum usually present; 41–54 (mean = 45.96 ± 3.49) dorsals in transverse row between dorsolateral crests at midbody; gular background in adult males light blue | E. microlepis |
– | White or cream spot posterior to tympanum absent; 37–47 (means = 41.63 ± 3.20 in E. azulae, 40.50 ± 1.90 in E. touzeti) dorsals in transverse row between dorsolateral crests at midbody; gular background in adult males cream or black | 18 |
18 | Vertebral scales in neck region in adult males similar in size to vertebrals in pelvic region; 45–57 (mean = 51.13 ± 4.05) gulars | E. azulae |
– | Vertebral scales in neck region in adult males more than twice as high as vertebrals in pelvic region; 42–48 (mean = 44.40 ± 2.22) gulars | E. touzeti |
For the type specimens of the species described in this paper we thank all collectors for their help in the field. Venegas is indebted to R. Wagter for logistic support in the field. We thank A. Varela and P. Santiana for editing the photographs; L.A. Coloma for providing photographs of the type specimens of E. altotambo; and C. Aguilar and J. Amanzo for providing valuable information. Specimens of the new species described in this paper were collected under collection permits 005-14-IC-FAU-DNB/MA issued by Ministerio del Ambiente, Ecuador, and N°08 C/C-2008-INRENA-IANP; 110-2007-INRENA-IFFS-DCB; 118-2007-INRENA-IFFS-DCB issued by Instituto Nacional de Recursos Naturales, Peru. This research was funded by The Systematics Association´s Systematics Research Fund (OTC), a Restricted Endowment Award from the Smithsonian Institution (KdQ, OTC), the Secretaría de Educación Superior, Ciencia, Tecnología e Innovación del Ecuador (OTC), Pontificia Universidad Católica del Ecuador (OTC), UCUMARI (PJV), and Gobierno Regional de San Martín GORESAM (PJV).
Specimens examined
Enyalioides cofanorum.—COLOMBIA: Amazonas: Puerto Nariño, 3°46'13"S, 70°22'59"W, 110 m, ICN 4229; ECUADOR: Orellana: 4km N Anangu on Garza Cocha at Hosteria La Selva, 260 m, MCZ 174428–29; Cononaco, QCAZ 5975; SPF, QCAZ 2710; transecto PBT, Pozo Capiron 2, QCAZ 7563; Tiputini Biodiversity Station, 0°37'5"S, 76°10'19"W, 215 m, QCAZ 8006; Yasuni National Park, bloque Shiripuno, 0°43'35"S, 76°43'36"W, QCAZ 3521; Sucumbíos: La Selva lodge, 0°24'0"S, 76°39'0"W, QCAZ 2935, 2961, 3951, 3953; Limoncocha, MCZ 157697; Santa Cecilia, 0°5'6"N, 76°59'33"W, 340 m, KU 105342 [paratype], 112180–81 [paratypes], 122118 [paratype], 146658 [holotype], 147584–85 [paratypes], 175308; Tarapoa, 0°7'60"S, 76°25'0"W, 283 m, FHGO 5764; Zamora Chinchipe: cuenca del rio Jamboe, Sakantza, 1230 m, FHGO 2342; PERU: Loreto: Ampiyacu river, Distrito Pevas, 3°19'0"S, 71°51'0"W, 100 m, CAS 8323.
Enyalioides heterolepis.—COLOMBIA: Antioquia: Dabeiba, río Amparradó, campamento Ingeominas, 6°42'0"N, 76°27'0"W, 805 m, ICN unassigned numbers (2 specimens); Municipio Frontino, corregimiento La Blanquita (Murrí), 800 m, IND-R 4229; Municipio Frontino, Vereda Venados, Parque Nacional Natural Las Orquídeas, afluente de la quebrada El Retiro, 6°33'0"N, 76°18'25"W, 850–950 m, ICN 9143; Cauca: bajo Calima, granja de la Secretaría de Fomento, 3°59'47"N, 76°58'28"W, ICN 4231; Guapí, 2°33'23"N, 77°51'50"W, ICN 4232, 4234–35; Gorgona Island, 2°58'31"N, 78°12'27"W, 30–120 m, FMNH 165387–88, ICN 824, 826–27, 832–38, 1045, 1247–53, 1325–26, 4237–44, 4521, 6515, ICN unassigned numbers (2 specimens), KU 192676–77; Guapí, on road to pipeline between Chansará-Cantadelicia, ICN 4233, 4236; Municipio de Junta, headwaters río Guapi, IND-R 3570; Quebrada Guangui, ICN unassigned number (1 specimen); Chocó: 5 km NW Playa de Oro, IND-R 3556; Bahía Solano, Parque Nacional Natural Utría, ICN unassigned number (1 specimen); headwaters of río San Juan, ca. 800 m, FMNH 165224; Quibdo, San josé de Purré, río Cabi, IND-R 5035; Serranía del Baudo, Alto del Buey, 6°6'0"N, 77°13'0"W, ICN 4245–46; Valle: Virology Field Station, USNM 151610–12; Valle del Cauca: 8 km W Danubio, río Anchicaya, KU 169853; Buenaventura, Base Naval Málaga, quebrada Valencia, 3°58'0"N, 77°18'0"W, ICN unassigned numbers (2 specimens); Dagua, Vereda La Elsa, 3°34'47"N, 76°46'54"W, 980 m, ICN 9091; km 6 on road Buenaventura-río Calima, 3°53'36"N, 77°4'11"W, 0 m, FMNH 165181–82; km 22 on road Buenaventura-río Calima, FMNH 165223; Municipio Restrepo, Vereda Alegre, Campo Chanco, 3°38'14"N, 76°13'44"W, 460 m, ICN 9093; río Raposo, above caserío El Tigre, 3°42'0"N, 77°5'60"W, 11 m, ICN 1501–02; no specific locality: ICN 9092, 9801, 11313; ECUADOR: El Oro: Gualtaco, USNM 211076; Esmeraldas: Alto Tambo, 253 m, QCAZ 5523; Bosque Protector La Chiquita, 30 km E San Lorenzo on road to Ibarra, QCAZ 3839; Corriente Grande, 70 m, QCAZ 3531; Jatun Sacha Field Station, Montañas Mache-Chindul, 41km W Quinindé, 0°21'21"N, 79°42'12"W, 600 m, FHGO 3200; Loma Linda, río Onzole, 95 m, QCAZ 3626; Mayronga, 100 m, QCAZ 2185–86, 2263–66; Reserva Ecológica Mache-Chindul, comunidad San Salvador, FHGO 4063; San Miguel de Cayapas, QCAZ 412; Los Ríos: Estación Biológica Río Palenque, 150-220 m, KU 146657, 164166, 180657–58, QCAZ 427, USNM 285451, 285454; Manabí: 38 km NW El Carmen, ca. El Carmen-Pedernales road, 330 m, KU 218380; Pichincha: 15 km NW La Florida, QCAZ 2844; La Perla, QCAZ 2025–26; Palma Real, USNM 211094; Puerto Quito, km 132 on road Calacalí-La Independencia, Hostería Selva Virgen, FHGO 4314; río Blanco, below mouth of río Toachi, USNM 211088; río Caoni, USNM 211095; río Toachi, between kms 100-110 on road to Santo Domingo de Los Colorados, USNM 211097–98; Santo Domingo de los Tsáchilas: km 30 Quinindé-Santo Domingo de los Colorados, USNM 211091; Santo Domingo de los Colorados, 600 m, KU 121090–91; PANAMA: Colón: Achiote, 40 m, KU 96688; Darién: Laguna, 820 m, KU 76050–53; ridge btw río Jaque & río Imamado, 730 m, KU 113490–94; SE slope Cerro Pirre, 1060 m, KU 96689–90; Tacarcuna, 550 m, KU 76047–48; San Blas: Armila, USNM 150121 Veragua (possibly Veraguas): MHNP 4067 [holotype].
Enyalioides laticeps.—BRAZIL: Fonteboa, upper Amazon, MHNP 6821 [holotype]; COLOMBIA: Amazonas: 50 km N Chorrera on Igará-Paraná, IND-R 1038–41; headwaters of río Caiwima, tributary of río Amacayacu, ca. 70 km NNE Puerto Nariño, MCZ 154482; Leticia, 4°12'55"S, 69°56'26"W, 83 m, ICN unassigned number (1 specimen); Parque Nacional Natural Amacayacu, cabaña Amacayacu, IND-R 4195; Parque Nacional Natural Amacayacu, río Amacayacu, Puerto Mogue, close to río Cabimas, IND-R 1037; Parque Nacional Natural Amacayacu, Mata-mata creek, 3 km W Mata-mata cabin, 3°41'0"S, 70°15'0"W, 150 m, ICN 9094; Puerto Rastrojo, río Miriti, IND-R 1920, 1929; río Amacayacu, tributary of río Amazonas, ca. 50km NNE Puerto Nariño, MCZ 156348; río Amacayacu-Caiwima, ca. 40km NNE Puerto Nariño, MCZ 154481; río Miriti Paraná, IND-R 1905; Caquetá: 30 km from mouth of río Cuemani, IND-R 1063–65; Florencia, MLS 117; Parque Nacional Natural Chiribiquete, río Mesay, Puerto Abeja, 0°5'27"N, 72°25'0"W, IAvH 4746; Guaviare: Chiribiquete, ICN unassigned number (1 specimen); Meta: río Guayabero, Angostura No.1, 2°17'0"N, 73°58'0"W, 300–350 m, ICN 1270; Cumaral, Vereda Juan Pablo II, 3°47'0"N, 73°55'0"W, ICN 7255; Guaguriba on road to Acacias, MCZ 156323; La Macarena, campamento Isama, río Duda, ICN 4230; La Macarena, Caño Guapayita, ICN 677; Las Salinas, 3 km NW Restrepo, 4°16'9"N, 73°35'9"W, 720 m, ICN unassigned number (1 specimen); La Macarena, río Duda, Parque Nacional Natural Los Tiniguas, campamento de primatología Puerto Chamuza, IND-R 4019–22, 4034; Serranía La Macarena, Caño Sardinata, 30 km W Vista Hermosa, IND-R 287; Villavicencio, 4°9'12"N, 73°38'6"W, 500 m, AMNH 35277, FMNH 30815, MLS 116; Villavicencio, Pozo Azul creek, 4°9'12"N, 73°38'6"W, 500 m, ICN 8341, ICN unsassigned numbers (2 specimens), MCZ 154334; Putumayo: ca. 10 km (airline) S Mocoa, 700–800 m, AMNH 106631; no specific locality: FMNH 165208, 165211; Vaupés: Caparú, surroundings of lake Taraira, 1°8'46"S, 69°29'14"W, ICN 8058–61; Estación Biológica Caparú, IND-R 4382; ECUADOR: Morona Santiago: cantón Taisha, parroquia Macuma, centro Shuar Macuma, 2°8'6.6"S, 77°42'54"W, 720 m, FHGO 5460; Arapicos, 1°51'0"S, 77°57'0"W, 981 m, USNM 211111; Napo: Ahuano, QCAZ 7014; Ávila, río Napo, CAS-SUR 8261–62; Tena, QCAZ 6054; Orellana: 7 km S río Tiputini, KU 299832–33; Estación Científica Yasuní, QCAZ 7388; Loreto, 0°40'0"S, 77°19'0"W, 451 m, USNM 211121; Parque Nacional Yasuni, Tambococha, FHGO 3692; Parque Nacional Yasuni, Tiputini, Ishpingo, FHGO 5346; Río Napo, Añangu, south bank, QCAZ 9503; Sacha Lodge, QCAZ 8884; Pastaza: Alto río, USNM 211146; Lorocachi, QCAZ 3222; Palmira, río Pastaza valley, AMNH 37554; río Capahuari, USNM 211122; río Huiyayacu, USNM 211128; río Pindo, USNM 211143; río Shiripuno, 1°5'0"S, 76°50'0"W, FHGO 1624; Sarayacu, USNM 211124; Villano, 1°30'0"S, 77°29'0"W, 388 m, QCAZ 8118, 8262; Sucumbíos: 2 km W Lago Agrio, KU 299835; Lago Agrio, KU 299834, KU 299836; río Cuyabeno, USNM 211113; Santa Cecilia, 0°3'0’’, N, 76°59'0’’, 340 m, KU 122104–05, 122110–11, 147931, 147939–41, 152497; San Jose, S Tarapoa, FHGO 4839; San Pablo de Kantesiaya, 0°15'0"S, 76°25'30"W, 240 m, FHGO 850; Zancudococha, 0°25'0"S, 75°30'0"W, 220 m, FHGO 304; PERU: Amazonas: Caterpiza, USNM 568575; Galilea, USNM 568576–80; Cusco: Pagoreni, río Camisea, 11°42'23"S, 72°54'11"W, 465m; Loreto: Explorama Lodge, jct río Yanamono & río Amazonas, KU 220493; Intuto, río Tigre, AMNH 60575; San Jacinto, 175 m, KU 222164; San Martin: San Martin, 14 km ESE Shapaja, 360 m, KU 212627; Ucayali: río Calleria, Colonia Calleria, 15 km from Ucayali, CAS 95143. NO SPECIFIC LOCALITY: ICN 1231.
Enyalioides oshaughnessyi.—ECUADOR: Esmeraldas: Bilsa Ecological Reserve, 225 m, QCAZ 6866; Guayas: cerro Masvale, QCAZ 9893; Los Ríos: Estacion Biológica Río Palenque, 150–220 m, KU 152597, USNM 285456–57, Estación Biológica Jauneche, 50 m, QCAZ 6899; Pichincha: Finca Victoria, 37 km SE Santo Domingo de los Colorados, MCZ 80958; Hotel Tinalandia, 15 km SE Santo Domingo de los Colorados, MCZ 145269; Puerto Quito, MCZ 164509; Recinto Playa Rica, on road Nanegal-Selva Alegre, QCAZ 7426; Silanchi, río Blanco, USNM 211102; Tandapi, MCZ 164789; Unión del Toachi, 300 m, QCAZ 5326, 6682; Santo Domingo de los Tsáchilas: 1 km N, 2 km E Santo Domingo de los Colorados, 620 m, KU 179417; 2 km E, 1 km S Santo Domingo de los Colorados, 600 m, KU 179416; Finca La Esperanza, 5 km W Santo Domingo de los Colorados, USNM 211105; Finca La Esperanza, 5 km W Santo Domingo de los Colorados, USNM 211106–07; Santo Domingo de los Colorados, KU 109630, USNM 211103, 211109. NO SPECIFIC LOCALITY: USNM 22448, 22450.