taxonID	type	description	language	source
285E87F7FFBFBC3A1DA0DD51C2C7FA5A.taxon	materials_examined	Type species: Echinosaura sulcarostrum Donnelly et al., 2006. LSID: urn: lsid: zoobank. org: act: A 7 C 758 C 4 - CD 77 - 4 CCF-BF 4 B- 79 C 1 D 1 A 1 C 854 Content: One species: Rheosaurus sulcarostrum (Donnelly et al., 2006) comb. nov. Diagnosis: Phenotypic characteristics of Rheosaurus include: (1) dorsal scales heterogeneous, with large or polygonal, longitudinally keeled scales intermixed with small irregular scales; (2) dorsal surface of the head with large, symmetrical scales; (3) internasal divided; (4) frontonasals three; (5) prefrontals paired; (6) frontal single; (7) frontoparietals paired; (8) interparietal well defined; (9) parietals paired; (10) rostral and mental striated; (11) three rows of scales between rostral and frontal scales; (12) postmental absent; (13) large chin shields in three pairs; (14) lower eyelid developed, with a palpebral disc divided into several, unpigmented scales; (15) ventral scales squared, not imbricated; (16) limbs pentadactyl, digits clawed; (17) femoral pores present in males (weakly developed) and absent in females. Rheosaurus differs from all other cercosaurines, including Echinosaura s. s. and its sister-group, in having striated rostral and mental scales and three rows of scales between rostral and frontal scales, and in lacking a postmental scale. It also differs from other cercosaurines, except Centrosaura, Echinosaura s. s., Gelanesaurus, Neusticurus and Potamites, in having heterogeneous dorsal scalation. For comparisons among related or similar genera see Figure 9 and Table 5. Etymology: Rheosaurus (gender masculine) is derived from the Greek ρέω, rheo (flow or stream) and σαύρα, saura (lizard), in reference to the riparian habit of this lizard. Geographic distribution: Pantepui bioregion of the Guiana Shield, from the vicinity of the Baramita area in north-western Guyana (100 m a. s. l.) to the Wokomung massif in west-central Guyana (Donnelly et al., 2006; Cole et al., 2013). Remarks: Donnelly et al. (2006) tentatively allocated sulcarostrum to Echinosaura and emphasized that this generic assignment reflected convenience rather than an understanding of the phylogenetic relationships of this species. They also anticipated that its disjunct distribution with respect to other Echinosaura and the differences in external head morphology, may reflect an independent phylogenetic history.	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFBFBC271EB7DDD6C4E0FD71.taxon	materials_examined	Type species: Echinosaura horrida Boulenger, 1890 (by original designation). Content: Seven species: Echinosaura brachycephala Köhler et al., 2004; E. centralis Dunn, 1944; E. horrida Boulenger, 1890; E. keyi (Fritts & Smith, 1969); E. orcesi Fritts, Almendáriz & Samec, 2002; E. palmeri Boulenger, 1911; E. panamensis Barbour, 1924. Diagnosis: Phenotypic characteristics of Echinosaura include: (1) dorsal scales heterogeneous, with small or granular, irregular scales intermixed with tubercular or spine-like scales; (2) dorsal surface of the head with large, symmetrical and paired scales anteriorly, and with irregular small scales posteriorly; (3) internasal single or divided; (4) frontonasals paired; (5) prefrontals usually absent; (6) frontal scale single or divided; (7) frontoparietals small, paired; (8) interparietal and postparietal scales not well defined (Fig. 10); (9) postmental large, single (reduced or absent in E. brachycephala); (10) chin shields usually large, in one pair (smaller in E. horrida, not differentiated in E. brachycephala) (Fig. 10); (11) lower eyelid developed, with a palpebral disc divided into several, unpigmented scales; (12) ventral scales squared (or rounded squares), not imbricated; (13) limbs pentadactyl, digits clawed; (14) males with well-developed femoral pores, females usually lacking them or having fewer femoral pores than males (if present, weakly developed or barely distinguishable). Morphological variation among species of Echinosaura is summarized in Table 2 and 3. Echinosaura differs from other cercosaurines, except Centrosaura, Gelanesaurus, Neusticurus, Potamites and Rheosaurus, in having heterogeneous dorsal scalation. It differs from the other genera with heterogeneous dorsal scalation in having small or granular, irregular dorsal scales intermixed with tubercular or spine-like scales (vs. small irregular or granular dorsal scales intermixed with large or polygonal keeled scales), several irregular scales on the parietal region (vs. regular scales on parietal region) and usually one pair of large chin shields (vs. usually more than one pair of large chin shields). A comparison of Echinosaura and closely related and morphologically similar genera is given in Table 5 and Figure 9. Etymology: Although not explicitly stated by Boulenger (1890), the name Echinosaura (gender feminine) is presumably formed from the Ancient Greek ἐχῖνος, echinos (hedgehog or urchin) and σαύρα, saura (lizard) – a spiny lizard. Geographic distribution: All species of Echinosaura are trans-Andean, extending from western Panama to north-western Ecuador. Most species occur in lowlands, but some reach as high as c. 2200 m a. s. l. Remarks: Our phylogenetic analysis corroborated the inclusion of the previously not-analysed species E. palmeri and E. panamensis in Echinosaura s. s., as recently hypothesized by Torres-Carvajal et al. (2016).	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA2BC271D92DAF3C5A0FBF4.taxon	description	(FIGS 6 A, 10 A)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA2BC271D92DAF3C5A0FBF4.taxon	diagnosis	Diagnosis: The following data are based on the original description and high-quality photographs of specimens deposited at QCAZ (Museo de Zoología, Pontificia Universidad Católica del Ecuador; photographs are available online through www. bioweb. bio). Echinosaura brachycephala possesses the following characteristics: (1) snout relatively short and rounded; (2) internasal single; (3) frontonasals paired [rarely with 1 – 4 small irregular scales between the frontonasals and the frontal (‘ prefrontals’)]; (4) frontal single (rarely divided); (5) frontoparietals paired; (6) supraoculars three (third supraocular smaller than first and second); (7) supralabials 3 – 5 (usually four); (8) infralabials three; (9) postmental reduced or absent; (10) chin shields not differentiated; (11) two paravertebral rows of tubercles or spine-like scales, slightly undulating, separated from each other by 4 – 6 small, irregular scales; (12) tubercular scales forming oblique lines on lateral surface of body; (13) ventral scales keeled, squared (or are rounded squares); (14) subdigital lamellae on the fourth finger 16 – 23; (15) subdigital lamellae on the fourth toe 23 – 32; (16) femoral pores per hind limb in males 7 – 9, in females 1 – 2; (17) dorsal and lateral surface of tail with small tubercles scattered along each caudal segment (coded as ‘ <pattern’); (18) subcaudals per caudal segment four (when tail is not regenerated); (19) dorsum brown or dark brown, relatively uniform, with some clear spots more or less visible; ventral surface of head and neck light brown with several scales bearing white spots, except on the edges near the mouth where there is a darker ground colour with some transverse cream blotches which extend to labial scales; ventral surface of body (including limbs and tail) mainly light brown spotted with dark brown and cream. Combination of these characteristics distinguishes Echinosaura brachycephala from its congeners. In addition, the distinctiveness of E. brachycephala and its sister-species, E. horrida (Fig. 2), is further corroborated by uncorrected pairwise distances of 18.8 % in ND 4 and 6.5 % in 16 S (Table 1). Geographic distribution: Echinosaura brachycephala is endemic to Ecuador. It occurs in the west versant of the Andes in Pichincha and Cotopaxi provinces at elevations between 690 and 1915 m a. s. l. (Fig. 8). Remarks: Most paratypes (42 specimens) were collected at the type locality (paratopotypes), whereas the remaining eight paratypes are from Tandapi, Pichincha province, Ecuador (Köhler et al., 2004). Although the distributions of Echinosaura brachycephala and E. horrida partially overlap, E. brachycephala seems to be more common above 1200 m a. s. l.	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA2BC231E87DC40C4E1FEEA.taxon	description	(FIGS 6 B, 7, 10 B, 11, 12)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA2BC231E87DC40C4E1FEEA.taxon	description	Neotype (designated here): MHUA-R 13332, an adult male (Figs 11, 12). Colombia, department of Antioquia, municipality of San Rafael, embalse Jaguas, Caño Girón (6 ° 23 ’ 54.672 ’ N, 75 ° 01 ’ 37.092 ’ W, 1275 m a. s. l.). Collected on 23 August 2017 by Juan D. Vásquez-Restrepo. Description of neotype: Adult male (Figs 11, 12), SVL = 60 mm, TRL = 27 mm, SL = 5.7 mm; HL = 15.2 mm, HW = 9.1 mm, TaL = 10.2 mm. Rostral scale single. Internasals two, longer than wide, with three longitudinal stretch marks, in contact with rostral anteriorly, nasals laterally, frontonasals posteriorly. Nasals subtriangular, in contact with internasals and first supralabials. Frontonasals about three times longer than wide, rectangular, with the posterior edges forming an obtuse angle and a depressed area in the middle part of each scale. Frontal single, wider anteriorly and thinner posteriorly, with a series of longitudinal, barely visible longitudinal ridges. Supraoculars three, first supraocular in contact with frontal and frontonasals, smaller than second and third; third supraocular in contact with three small, keeled ciliaries. Lower eyelid with a palpebral disc divided into three large, unpigmented scales. A large polygonal scale in loreal region not in contact with the supralabials, except on the right side where it is fused with the first large subocular scale. Loreal in contact with preciliary, frontonasal and nasal. Suboculars in four small rows, in contact with a fifth row formed by five large scales on the left side and four on the right side (fifth scale fused with loreal), which are in contact with second to fifth supralabials. Frontoparietals paired, small, well defined. Parietal region covered with small irregular scales. Five upper and four lower labials on both sides. Mental posteriorly in contact with postmental. Postmental trapezoidal, in contact with first infralabial and anterior half of second infralabial. Chin shields large, in one pair, about three times longer than wide, in contact with posterior half of the second and anterior half of the third infralabials, separated from infralabials by three rows of small, hexagonal scales arranged longitudinally, larger than adjacent. Gular region with small, semicircular and tubercular scales, becoming spine-like scales that extend to posterior part of head in dorsal view in four longitudinal rows. Two of these lines are continuous and parallel, extending on paravertebral region and separated from each other by 6 – 7 irregular, small scales. The remaining two rows extend on upper coastal region, also formed by spine-like scales, but forming a discontinuous longitudinal and sinuous line. On the sides, there is a series of spine-like scales forming oblique lines. Spine-like scales on dorsum separated from each other by small, irregular or tubercular scales. Limbs pentadactyl, digits clawed. Subdigital lamellae on the fourth finger 15 / 14 (second one on left side divided); subdigital lamellae on the fourth toe 21. Scales on pectoral region flat, rounded, becoming six rows of squared and keeled scales on the belly. Tail with two parallel ridges dorsally, formed by a series of increasing in size scales from the anterior margin of each caudal segment, with more-developed scales forming longitudinal ridges laterally. Tail complete, with three subcaudal scales per caudal segment. Dorsum light brown, with some dark-brown scales on head, barely yellowish blotches on body and limbs, and two cream spots at the base of the tail. Scales of ventral surface of head dark brown (almost black), most of them with a cream spot on the centre. Mental and labials with whitish colouration on the posterior edges. Suture between postmental and chin shields with a white transverse stripe. Scales on belly with an irregular pattern of chessboard (black and cream scales). Ventral surface of tail dark brown (almost black) from the basis, except by a few whitish scales on the anterior and posterior part. Diagnosis: The following data are based on 70 examined specimens (referred material listed in Supporting Information, Appendix SII). Echinosaura centralis possesses the following characteristics: (1) snout pointed; (2) internasal divided longitudinally (rarely single); (3) frontonasals paired; (4) frontal single and long (it extends up to the middle of the eyes); (5) frontoparietals paired; (6) supraoculars two or three, large; (7) supralabials 4 – 6 (usually five); (8) infralabials 3 – 5 (usually four); (9) postmental single; (10) large chin shields in one pair; (11) two paravertebral ridges, separated from each other by usually five or more small, irregular scales; (12) spine-like scales forming oblique lines on lateral surface of body; (13) ventral scales squared, usually keeled; (14) subdigital lamellae on the fourth finger 12 – 18; (15) subdigital lamellae on the fourth toe 20 – 27; (16) femoral pores per hind limb in males 4 – 9; (17) dorsal surface of tail with two parallel ridges, formed by a series of increasing in size scales from the anterior margin of each caudal segment; lateral surface of tail with more-developed scales forming longitudinal ridges (coded as ‘ = pattern’); (18) subcaudals per caudal segment three (when tail is not regenerated); (19) dorsum brown or dark brown, relatively uniform, sometimes with yellowish marks and / or two spots more or less distinguishable at the base of tail; ventral surface of head and neck mainly light brown, except on the anterior part where there is a darker ground colour with some transverse cream blotches, extending to labial scales; ventral surface of body (including limbs) mainly light brown spotted with dark brown or cream, while that of tail almost completely dark. Colouration in preservative is darker in some specimens. Combination of these characteristics distinguishes Echinosaura centralis from its congeners. In addition, the distinctiveness of E. centralis and its sister-species E. palmeri (Fig. 2) is corroborated by uncorrected pairwise distances of 17.6 % in ND 4 and 5.6 % in 16 S (Table 1). Geographic distribution: Echinosaura centralis is endemic to Colombia. It occurs in the Cordillera Central from Tolima to Bolívar departments, the adjacent Magdalena Valley and the west versant of the Cordillera Oriental in Cundinamarca and Boyacá departments. Most records are above 1000 m a. s. l., but its elevation ranges from 50 to 2200 m a. s. l. (Fig. 8). Natural history: We have observed individuals of Echinosaura centralis diving into the streams and swimming short distances after being disturbed. Variation: The referred specimens are similar to the neotype with the following noteworthy exception: internasal scale single (an unusual condition) in MHUA-R 10132, 11245, 11440, 12111, 12273 and 13212.	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA6BC231DD7D965C2FBFC3C.taxon	description	(FIG 6 C, 10 C)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA6BC231DD7D965C2FBFC3C.taxon	materials_examined	Type material: Boulenger (1890: 83) described Echinosaura horrida based on two specimens deposited in the lizard collection of the British Museum (BMNH), a female and a young, neither of which was referenced with a specific collection number. Boulenger also did not designate a holotype. However, according to the data portal of the BMNH (http: // data. nhm. ac. uk) and Torres-Carvajal (2001), there are two type specimens (syntypes following the Art. 73.2 of the ICZN) of E. horrida (BMNH 1946.8.31.60 and 1946.8.31.61) whose information matches the original description. Diagnosis: The following data are based on 13 examined specimens (referred material listed in Supporting Information, Appendix SII) and data taken from the literature (Uzzell, 1965; Köhler et al., 2004). Echinosaura horrida possesses the following characteristics: (1) snout pointed, relatively short; (2) internasal single; (3) frontonasals paired; (4) frontal single, short; (5) frontoparietals paired; (6) supraoculars two or three, large; (7) supralabials usually five or six; (8) infralabials three (some specimens from Ecuador have four supralabials and / or infralabials); (9) postmental single; (10) chin shields short, rounded and in one pair; (11) two longitudinal, parallel and juxtaposed paravertebral ridges; (12) series of spine-like scales forming oblique lines on lateral surface of body; (13) ventral scales squared, keeled; (14) subdigital lamellae on the fourth finger 14 – 19; (15) subdigital lamellae on the fourth toe 21 – 25; (16) femoral pores per hind limb in males usually 7 – 10; (17) dorsal surface of tail with two parallel ridges, formed by a series of scales increasing in size from the anterior margin of each caudal segment; lateral surface of tail with more-developed scales forming longitudinal ridges (coded as ‘ = pattern’); (18) subcaudals per caudal segment three (when tail is not regenerated); (19) dorsum brown, sometimes with faint yellowish spots; ventral surface of head and neck mostly light brown, except on the anterior part where there is a darker ground colour, with some transverse cream blotches extending to labial scales; ventral surface of body (including limbs) mainly light brown spotted with dark brown and cream, while that of tail has blotches or dark transverse marks. Colouration in preservative similar to that in life. Combination of these characteristics distinguishes Echinosaura horrida from its congeners. In addition, the distinctiveness of E. horrida and its sister-species E. brachycephala (Fig. 2) is further corroborated by uncorrected pairwise distances of 18.8 % in ND 4 and 6.5 % in 16 S (Table 1). Geographic distribution: Echinosaura horrida occurs on the Pacific versant of Ecuador and Colombia, from central-west Ecuador in the western part of Cotopaxi province to Isla Gorgona in Colombia and the adjacent mainland. This is mainly a lowland species, but its elevation ranges from 10 to c. 1546 m a. s. l. (Fig. 8). Remarks: Echinosaura horrida occurs in sympatry with E. keyi in Ecuador, in the vicinity of Mataje and in El Placer, Esmeraldas province and in La Florida, Pichincha province (Fritts et al., 2002).	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA6BC221E67DC2FC474FB36.taxon	description	(FIG. 6 D)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA6BC221E67DC2FC474FB36.taxon	diagnosis	Diagnosis: The following data are based on the original description, literature (Köhler et al., 2004) and high-quality photographs of specimens deposited at QCAZ (available online through www. bioweb. bio). Echinosaura keyi possesses the following characteristics: (1) snout relatively short; (2) internasal longitudinally divided; (3) frontonasals paired; (4) frontal extending up to the middle of the eyes, usually single but could be divided transversely; (5) frontoparietals paired; (6) supraoculars three (third supraocular smaller than first and second); (7) supralabials four; (8) infralabials 3 – 4; (9) tympanum covered with small, granular scales; (10) postmental single, short; (11) large chin shields in one pair; (12) two longitudinal paravertebral ridges formed by spine-like scales, discontinuous on posterior portion of body; (13) series of conical scales of different sizes forming oblique lines on lateral surface of body; (14) ventral scales squared, smooth; (15) femoral pores per hind limb in males usually 8 – 11; (16) tail with six longitudinal rows of enlarged conical scales on the posterior part of each caudal segment (coded as ‘ ^ pattern’); (17) subcaudals per caudal segment three (when tail is not regenerated); (18) dorsum dark brown or almost black, with some yellow or cream reticulate spots; ventral surface of head and neck mainly cream with some dark spots; ventral surface of body (including limbs) mainly cream or light brown spotted with dark brown, while that of tail spotted with light and dark brown. Combination of these characteristics distinguishes E. keyi from its congeners. In addition, the distinctiveness of E. keyi and its congeners is corroborated by a minimal uncorrected pairwise distances of 17.2 % in ND 4 and 6.7 % in 16 S (Table 1). Geographic distribution: Echinosaura keyi is endemic to Ecuador. Its known distribution is restricted to the Pacific versant of north-western Ecuador, including Santo Domingo, Pichincha, Esmeraldas, Imbabura and Carchi provinces. Its elevational range is from 200 to 1235 m a. s. l. (Fig. 8). Remarks: Echinosaura keyi occurs in sympatry with E. horrida in the vicinity of Mataje and in El Placer, Esmeraldas province and in La Florida, Pichincha province (Fritts et al., 2002). It is possible that this species extends into southern Colombia.	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA7BC221DE2DD32C38AFA69.taxon	description	(FIGS 6 E, 10 D)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA7BC221DE2DD32C38AFA69.taxon	diagnosis	Diagnosis: The following data are based on nine examined specimens (referred material listed in Supporting Information, Appendix SII) and the original description. Echinosaura orcesi possesses the following characteristics: (1) snout pointed, long; (2) internasal single; (3) frontonasals paired; (4) frontal long, single; (5) frontoparietals paired; (6) supraoculars large, usually two or three; (7) supralabials 3 – 5 (usually three); (8) infralabials 2 – 3 (usually three); (9) postmental single; (10) large chin shields in one pair; (11) dorsally two discontinuous sinuous or S-shaped ridges formed by spine-like scales; (12) alternate tubercular scales on lateral surface of body; (13) ventral scales smooth, squared (or are rounded squares); (14) subdigital lamellae on the fourth finger 20 – 22; (15) subdigital lamellae on the fourth toe 30 – 36; (16) femoral pores per hind limb in males usually 9 – 16; (17) each caudal segment without crests, with a pair of more-developed scales or tubercles on the distal region forming transverse rings (coded as ‘: pattern’); (18) subcaudals per caudal segment five or six (when tail is not regenerated); (19) dorsum brown with transverse alternate darker bands, and with lateral ocelli more or less visible; ventral surface of head and neck cream or clear brown, except on the anterior part where there are some transverse dark brown blotches that extend to labial scales; ventral surface of body (including limbs and tail) mainly cream or reddish light brown spotted with dark (in some cases dark spotted with cream). Colouration in preservative darker. Combination of these characteristics distinguishes E. orcesi from its congeners. In addition, the distinctiveness of E. orcesi and its congeners is corroborated by a minimal uncorrected pairwise distances of 17.4 % in ND 4 and 7.4 % in 16 S (Table 1). Geographic distribution: Echinosaura orcesi occurs on the western versant of the Cordillera Occidental and in the Pacific region of Colombia, from Anchicayá region in Valle del Cauca department to north-western Ecuador in Esmeraldas and Carchi provinces. Its elevational range extends from 50 to c. 866 m a. s. l. (Fig. 8). Remarks: Echinosura orcesi occurs in sympatry with E. palmeri on the Anchicayá drainage in the Valle del Cauca department, south-western Colombia. In Ecuador, near the border with Colombia in Esmeraldas and Carchi provinces, E. orcesi overlaps in its distribution range with E. horrida and E. keyi. Köhler et al. (2004) also reported that these three species occur sympatrically in Paramba, Imbabura province.	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA7BC211E7ADE18C269FD99.taxon	description	(FIGS 6 G, 6 H, 7, 10 F)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA7BC211E7ADE18C269FD99.taxon	materials_examined	Type material: Boulenger (1911) described Echinosaura palmeri based on a single male deposited in the lizard collection of the British Museum, which he did not designate explicitly as the holotype. However, according to Art. 73.1.2 of the ICZN (1999), this specimen is assumed to be the holotype. Boulenger also did not provide a museum number for it. However, according to the data portal of the BMNH (http: // data. nhm. ac. uk), there is a type specimen of E. palmeri (BMNH 1946.8.31.24) whose information matches the original description. Thus, following the Art. 72.4.1.1 of the ICZN, we consider this specimen as the holotype of E. palmeri. Diagnosis: The following data are based on 70 examined specimens (referred material listed in Supporting Information, Appendix SII). Echinosaura palmeri possesses the following characteristics: (1) snout pointed; (2) internasal divided longitudinally; (3) frontonasals paired; (4) frontal long, single, extending up to the middle of the eyes; (5) frontoparietals paired; (6) supraoculars usually two or three; (7) supralabials 4 – 5 (usually five); (8) infralabials 4 – 5 (usually four); (9) postmental single; (10) large chin shields in one pair; (11) two longitudinal, parallel paravertebral ridges, separated from each other by 3 – 8 (usually five) small irregular scales; (12) series of spine-like scales forming oblique lines on lateral surface of body; (13) ventral scales squared, usually smooth (rarely weakly keeled); (14) subdigital lamellae on the fourth finger 13 – 16; (15) subdigital lamellae on the fourth toe 19 – 25; (16) femoral pores per hind limb in males usually 7 – 8; (17) each caudal segment usually without crests, with a more-developed scales or tubercles on the distal region forming transverse rings (coded as ‘: pattern’), but in some cases dorsally with some small scales oblique forming a V shape (coded as ‘ <pattern’); laterally without more-developed scales forming longitudinal ridges; (18) subcaudals per caudal segment three (when tail is not regenerated); (19) dorsum brown or dark brown relatively uniform, sometimes with yellowish marks and / or two spots more or less distinguishable at the base of the tail; ventral surface of head and neck mainly light brown, except in the anterior part where there are some transverse dark brown blotches that extend to labial scales; ventral surface of body (including limbs) mainly cream or light brown spotted with dark brown, while that of tail almost completely dark. Colouration in preservative could be darker. Combination of these characteristics distinguishes Echinosaura palmeri from its congeners. In addition, the distinctiveness of E. palmeri and its sister-species, E. centralis (Fig. 2), is further corroborated by uncorrected pairwise distances of 17.9 % in ND 4 and 5.6 % in 16 S (Table 1). Geographic distribution: Echinosaura palmeri occurs in the southern part of Valle del Cauca department in Colombia, through the Pacific and adjacent Cordillera Occidental, to the Darien region in Panama. This is mainly a lowland species (Uzzell, 1965; Köhler et al., 2004), but its range extends from 2 to c. 1520 m a. s. l. (Fig. 8; Jaramillo et al., 2010). Remarks: Echinosaura palmeri occurs in sympatry with E. orcesi in the Anchicayá region in Colombia, near the southern distribution limit of E. palmeri and the northern one of E. orcesi.	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA4BC201E65DA85C3BAFDDC.taxon	description	(FIGS 6 F, 10 E)	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
285E87F7FFA4BC201E65DA85C3BAFDDC.taxon	materials_examined	Type material: Barbour (1924) described Echinosaura panamensis based on a single specimen (MCZ 17746), although he mentioned a series of ‘ no less than five [specimens] ’. However, Barbour did not provide museum numbers for the remaining specimens (paratypes according to the Art. 74.4.5 of the ICZN). Later, Uzzell (1965) mentioned the existence only of three paratypes (MCZ 18857 – 59), but according to the written museum catalogue (available online), there is a fourth specimen (MCZ 18860). Additionally, in the MCZ data portal (http: // mczbase. mcz. harvard. edu, accessed July 30 of 2019) this last specimen, also flagged as paratype, was sent to Chicago in 2015. Its current location is unknown. Although the existence of a fourth paratype is not in conflict with the original description (i. e. at least five), we consider as paratypes only those revised by Uzzell (1965), until physical evidence of the existence of MCZ 18860 is given. The above follows Art. 72.4.1.1 of the ICZN. Diagnosis: The following data are based on the original description, eight examined specimens (referred material listed in Supporting Information, Appendix SII) and data taken from the literature (Uzzell, 1965; Köhler et al., 2004). Echinosaura panamensis possesses the following characteristics: (1) snout pointed; (2) internasal divided longitudinally; (3) frontonasals paired; (4) frontal divided transversely, the anterior being larger than the posterior; (5) frontoparietals paired; (6) supraoculars large, usually three; (7) supralabials 4 – 6 (usually five); (8) infralabials 4 – 6 (usually five); (9) postmental single; (10) large chin shields in one pair; (11) two longitudinal paravertebral ridges formed by tubercular scales arranged in zig-zag or sinuous pattern; (12) series of tubercular scales forming oblique lines on lateral surface of body; (13) ventral scales squared, smooth; (14) subdigital lamellae on the fourth finger 13 – 16; (15) subdigital lamellae on the fourth toe 20 – 22; (16) femoral pores per hind limb in males usually 3 – 9; (17) each caudal autotomic segment dorsally with two small crests, oblique or V-shaped (coded as ‘ <pattern’) and sometimes almost straight; laterally, last scales of each caudal segment forming transverse rings, and without more-developed scales forming longitudinal ridges; (18) subcaudals per caudal segment three (when tail is not regenerated); (19) dorsum brown, sometimes with yellowish marks	en	Vásquez-Restrepo, Juan D., Ibáñez, Roberto, Sánchez-Pacheco, Santiago J., Daza, Juan M. (2020): Phylogeny, taxonomy and distribution of the Neotropical lizard genus Echinosaura (Squamata: Gymnophthalmidae), with the recognition of two new genera in Cercosaurinae. Zoological Journal of the Linnean Society 189: 287-314
