A new species of cat-eyed snake (Serpentes: Dipsadinae: Leptodeirini) from the Andes of southern Ecuador Author Torres-Carvajal, Omar 0000-0003-0041-9250 Museo de Zoología, Departamento de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Apartado 17 - 01 - 2184, Quito, Ecuador. & omartorcar @ gmail. com; https: // orcid. org / 0000 - 0003 - 0041 - 9250 & omartorcar @ gmail. com; https: // orcid. org / 0000 - 0003 - 0041 - 9250 omartorcar@gmail.com Author Sánchez-Nivicela, Juan C. 0000-0001-7971-1216 Instituto de Ciencias Naturales, Universidad Nacional de Colombia, A. A. 7495, Bogotá, Colombia. & juan. sanchezn 13 @ gmail. com; https: // orcid. org / 0000 - 0001 - 7971 - 1216 & Instituto Nacional de Biodiversidad, calle Rumipamba 341 y Av. de los Shyris, Casilla 17 - 07 - 8976, Quito, Ecuador. & juan. sanchezn 13 @ gmail. com; https: // orcid. org / 0000 - 0001 - 7971 - 1216 & valeposse _ 92 @ hotmail. com; https: // orcid. org / 0000 - 0003 - 0199 - 7554 juan.sanchezn13@gmail.com Author Posse, Valentina 0000-0001-7971-1216 Instituto Nacional de Biodiversidad, calle Rumipamba 341 y Av. de los Shyris, Casilla 17 - 07 - 8976, Quito, Ecuador. & juan. sanchezn 13 @ gmail. com; https: // orcid. org / 0000 - 0001 - 7971 - 1216 & valeposse _ 92 @ hotmail. com; https: // orcid. org / 0000 - 0003 - 0199 - 7554 & Laboratorio de Herpetología, Museo de Zoología, Escuela de Biología, Ecología y Gestión, Universidad del Azuay, Avenida 24 de mayo 7 - 77 y Hernán Malo, Apartado 01 - 01 - 981, Cuenca, Ecuador. & valeposse _ 92 @ hotmail. com; https: // orcid. org / 0000 - 0003 - 0199 - 7554 & elvisceli @ hotmail. com; https: // orcid. org / 0000 - 0001 - 9825 - 7477 juan.sanchezn13@gmail.com Author Celi, Elvis 0000-0003-0199-7554 Laboratorio de Herpetología, Museo de Zoología, Escuela de Biología, Ecología y Gestión, Universidad del Azuay, Avenida 24 de mayo 7 - 77 y Hernán Malo, Apartado 01 - 01 - 981, Cuenca, Ecuador. & valeposse _ 92 @ hotmail. com; https: // orcid. org / 0000 - 0003 - 0199 - 7554 & elvisceli @ hotmail. com; https: // orcid. org / 0000 - 0001 - 9825 - 7477 valeposse_92@hotmail.com Author Koch, Claudia 0000-0002-7115-2816 Department of Herpetology, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Adenauerallee 160, 53113 Bonn, Germany. & C. Koch @ leibniz-zfmk. de; https: // orcid. org / 0000 - 0002 - 7115 - 2816 och@leibniz-zfmk.de text Zootaxa 2020 2020-12-16 4895 3 357 380 journal article 9251 10.11646/zootaxa.4895.3.3 58efa18a-128f-4c5d-897f-99e5c13b660e 1175-5326 4326719 1AC38644-0F08-4127-AFE0-BDFDD6B0A2E2 Leptodeira misinawui sp. nov. ( Figs. 2–3 ) Holotype . An adult male, MZUA-RE 0253 , collected on 22 April 2015 by Elvis Celi in Poetate (- 3.417S , - 79.227W , 2,734 m above sea level ; hereafter asl), cantón Susudel , Azuay province , Ecuador . Paratypes (N=3). ECUADOR : Azuay : adult female, MZUA-RE 0252 , with the same collection data as the holotype; adult male, QCAZ 16510 , collected on 18 March 2018 by Diego Almeida on the road Santa Isabel-Pasaje (- 3.30S , - 79.340W , 971 m asl ) ; juvenile, QCAZ 16511 , collected on 20 March 2018 by Diego Almeida on the road Santa Isabel-Pasaje (- 3.320S , - 79.353W , 950 m asl ) . FIGURE 2. Male holotype (MZUA-RE 0253) and female paratype (MZUA-RE 0252) of Leptodeira misinawui sp. nov. in preservative. Dorsal (A) and ventral (B) views of body, and dorsal, ventral and lateral views of head (C) of the holotype. Dorsal (D) and ventral (E) views of body, and dorsal, ventral and lateral views of head (F) of the paratype. Photographs by J.C. Sánchez. Scale bar (C, F) = 10 mm. FIGURE 3. General view of Leptodeira misinawui sp. nov. in life. Top: holotype (MZUA-RE 0253); bottom: paratype (QCAZ 16510), adult male from Azuay province, road Santa Isabel-Pasaje. Photographs by E. Celi (top) and G. Pazmiño (bottom). Diagnosis. Leptodeira misinawui can be distinguished from other congeners by unique combination of the following characters: 19 (25%) or 21 (75%) midbody dorsal scale rows, reducing to 16 (25%) or 17 (75%) posteriorly; 2–3 preoculars (usually 2); vertebral and paravertebral rows not enlarged; single nape bar; faint intercalary spots; and 54–72 (average 63.25 ± 7.89 SD) body blotches rarely forming a continuous zig-zag stripe, 2–3 dorsal scales in length, half the length of light interspaces, and extending laterally onto scale rows 7–8. The new species differs from Leptodeira ornata (in parentheses) from western Ecuador in having 54–72 dorsal blotches along body (vs. 30–47), and tail length/body length 0.30 in females, 0.36–0.39 in males (vs. 0.21–0.26 in females, 0.23–0.30 in males). Similarly, L. misinawui has more dorsal blotches along body than L. larcorum (42–54). The dorsal blotches of L. misinawui are noticeably smaller (usually <2 scales long and <5 scales wide) than those of L. ornata and L. larcorum (usually> 2 scales long and> 10 scales wide). Although similar in morphology, the hemipenis of L. misinawui differs from those of L. larcorum and L. ornata in having shorter spines, with an average spine length/spine base-width ratio of 1.6 (vs. 3.3 and 2.2, respectively). A comparison of scutellation and size between Leptodeira misinawui , L. ornata from western Ecuador , and L. larcorum is presented in Table 5 . Description of the holotype . Male, SVL= 445 mm , TL= 173 mm . Head distinct from neck; rostral barely visible from above; head two times longer than broad and almost 2.4 times longer than deep. Eye slightly protuberant, occupying 16.5% of length of head, with elliptical pupil visible from anterior, lateral, dorsal, and ventral aspects; rostral 1.8 times wider than high, concave in anterior view, and narrowly visible from above. Prefrontals hexagonal, slightly longer than broad, twice the length of internasals, each in contact with its mate and with frontal posteriorly, supraocular (left only) posterolaterally, preocular posterolaterally, nasal anterolaterally, internasal anteriorly, and loreal laterally; frontal approximately pentagonal, 1.3 times longer than wide (greatest width), and its length about the same distance from its anterior edge to tip of snout. Supraocular longer than wide, in contact with frontal medially, preocular anteriorly, eye laterally, postocular posterolaterally, and parietal posteriorly; frontal slightly shorter than suture between parietals; broad parietals, about 2 times longer than wide; interparietal suture 1.1 times shorter than distance from frontal to tip of snout; nasal plate single, centrally pierced by large naris ( 1.17 mm in diameter), in contact with rostral anteriorly, internasal dorsally, prefrontal posterodorsally, loreal posteriorly, and first and second supralabials ventrally; preoculars two, the ventral one hexagonal and much smaller than the dorsalmost; postoculars two (an extra tiny scale between postoculars on left side), the lower less than half the size of the upper one; temporals 1+2 (the upper second temporal broken into two small scales on left side); supralabials eight, first and second in contact with nasal, second and third in contact with loreal, third and fourth in contact with preocular, fourth and fifth bordering the orbit; infralabials 10, with first to fifth in contact with anterior genial, and fifth and sixth touching posterior genial; first pair of infralabials in contact medially behind mental; genials in two pairs of similar length ( Fig. 2C ). Body higher than wide, rounded ventrolaterally; dorsal scales 19–21–17, smooth, juxtaposed or subimbricate, with paired apical pits; scales of vertebral row similar in size to adjacent rows; ventrals 180; anal plate divided; subcaudals divided, 90 ( Fig. 2A,B ). FIGURE 4. Hemipenes of Leptodeira misinawui sp. nov. (top, MZUA-RE 0253, holotype), L. ornata (middle, MZUA-RE 0173), and L. larcorum (bottom, ZFMK 91794). Views from left to right: lateral, asulcate, and sulcate. Photographs by J.C. Sánchez and M. Flecks. Scale bar = 5 mm. TABLE 4 . Uncorrected p-distances between 15 species of Leptodeira for the ND4 gene region. Taxonomy is based on the phylogenetic tree in Figure 1.

L . annulata	L. ashmeadii	L . maculata	L . rhombifera	L. bakeri	L. frenata	L . larcorum	L . nigrofasciata L . polysticta	L. punctata	L. rubricata	L. ornata	L. misinawui	L . splendida bressoni	L. s. splendida
L. annulata
L. ashmeadii	0.08
L. maculata	0.10	0.09
L. rhombifera	0.10	0.08	0.07
L. bakeri	0.08	0.03	0.08	0.09
L. frenata	0.15	0.14	0.15	0.13	0.14
L. larcorum	0.07	0.04	0.09	0.09	0.04	0.13
L. nigrofasciata	0.17	0.16	0.17	0.15	0.18	0.17	0.17
L. polysticta	0.10	0.08	0.10	0.10	0.09	0.14	0.08	0.16
L. punctata	0.15	0.15	0.15	0.15	0.14	0.15	0.14	0.18 0.14
L. rubricata	0.10	0.09	0.07	0.01	0.09	0.13	0.09	0.15 0.10	0.15
L. ornata	0.07	0.04	0.08	0.08	0.05	0.14	0.04	0.17 0.08	0.13	0.08
L. misinawui	0.06	0.05	0.08	0.07	0.06	0.13	0.04	0.16 0.08	0.14	0.08	0.05
L. splendida bressoni	0.13	0.13	0.14	0.13	0.14	0.13	0.13	0.19 0.13	0.14	0.13	0.11	0.12
L. s. splendida	0.14	0.13	0.14	0.13	0.14	0.15	0.13	0.18 0.14	0.15	0.13	0.12	0.13	0.03
L. uribei	0.15	0.13	0.14	0.14	0.14	0.01	0.13	0.17 0.13	0.14	0.14	0.14	0.13	0.14	0.15

Dorsal background light brown, with a longitudinal series (~67 on body and ~38 on tail) of dark brown paravertebral blotches on each side, some of which are fused with the corresponding opposite blotch, but never forming a zig-zag line. Paravertebral blotches on body 2–3 dorsal scales in length, half the length of dorsal light interspaces, extending laterally onto scale rows 7–8, and becoming smaller towards the tip of the tail. Faint intercalary spots on dorsal row 4, or between rows 4 and 5. Ventral surface of body cream with minute dark brown dots, more numerous at lateral ends of ventral scales, head (ventrally), and margins of subcaudals. Dark brown postocular stripe from eye to corner of mouth, extending from postoculars across ventral edges of primary and secondary temporals, dorsal edge of supralabial VI, upper half of supralabial VII, and anterior half of supralabial VIII; another dark stripe from nostril across loreal and upper preocular to orbit. Infralabials cream, densely marked with tiny dark dots. Head background same as dorsum, with irregular dark brown marks that are denser on parietals. Longitudinal nape bar extending from posterior margins of parietals to about third middorsal row; two short longitudinally elongated blotches on nape, the left one fused with nape bar. Iris bronze ( Fig. 3 ). Fully everted hemipenis ( Fig. 4 ) longer ( 11.3 mm ) than wide ( 5.5 mm ), extending to fifth/sixth subcaudal in situ, unilobed, unicapitate; capitulum rounded and finely crenulate, with a deep fold at the base; hemipenial body with small spinules at its base and large spines distally, which are more numerous laterally; sulcus spermaticus unforked. Variation. Intraspecific variation of Leptodeira misinawui in scutellational and meristic characters is presented in Table 5 . Paratypes are generally similar to holotype in color patterns. Paratype MZUA-RE 0252 has less elongated blotches on nape (circular right, ovoid left) and the nape stripe is not fused to the first body blotches ( Fig. 2 ). In paratype QCAZ 16511, the longitudinal nape bar is fused to the right nape blotch, whereas in QCAZ 16510 both nape blotches are fused to the nape bar forming a butterfly-shaped mark ( Fig. 3 ). The two examined males have more subcaudals and a longer tail than the female ( Table 5 ). Etymology. The specific epithet “ misinawui ” derives from the Quechua-Cañari words “ misi ” meaning cat, and “ ñawui ” meaning eye, and refers to the similarity of the eye of the new species to the eye of a cat. This epithet honors the Quechua-Cañari culture from southern Ecuador . We propose the Standard English Name: Jubones Cat-Eyed Snakes and the Standard Spanish Name: Serpientes Ojo de Gato del Jubones. Distribution and natural history. Leptodeira misinawui is known from upper basin of Jubones River, Azuay Province , at elevations between 950–2,734 m asl ( Fig. 5 ). This area corresponds to Evergreen Montane Forest of the southern Ecuadorian Andes and is characterized by a series of small isolated mountains within an inter-Andean valley (Ministerio del Ambiente del Ecuador 2012). Habitat at the type locality is characterized by medium size shrubs < 2m high, with scattered herbaceous vegetation that usually accumulate water ( Fig. 6 ). Individuals of L. misinawui were encountered active at night near roots of shrubs, near trails, and in a small water pool. FIGURE 5 . Known distribution of Leptodeira misinawui (diamonds), L. ornata in western Ecuador (circles), and L. larcorum (triangles). Filled symbols represent specimens sequenced in this study. FIGURE 6 . Landscape view of Jubones River Basin in southern Ecuador (A), and close-up of the type-locality habitat of Leptodeira misinawui (B). Photographs by S.R. Ron (A) and E. Celi (B). TABLE 5 . Selected meristic and morphometric characters for Leptodeira misinawui , L. ornata from western Ecuador, and L. larcorum from Peru. Range (first line) and mean ± SD (second line) are presented when appropriate.

Character	L. misinawui ECUADOR N=4	L. ornata ECUADOR N=61	L. larcorum PERU N= 21
Dorsal scale rows on neck	19 (75%) 21 (25%)	17 (1.7%) 19 (53.3%) 20 (6.6%) 21 (33.3%) 22 (3.4%) 23 (1.7%)	(N=20) 15 (5%) 17 (5%) 19 (60%) 20 (30%)
Dorsal scale rows at midbody	21 (75%) 19 (25%)	19 (13.1%) 21 (65.6%) 22 (1.6%) 23 (19.7%)	(N=20) 17 (5%) 19 (10%) 20 (25%) 21 (60%)
Dorsal scale rows anterior to cloaca	17 (75%) 16 (25%)	13 (1.6%) 14 (1.6%) 15 (54.2%) 16 (16.4%) 17 (26.2%)	(N=20) 14 (5%) 15 (60%) 16 (15%) 17 (15%) 18 (5%)
Ventrals in males	180–181 (N=2) 180.5 ± 0.71	176–197 (N=37) 187.49 ± 5.19	186–196 (N=10) 189 ± 3.56
Ventrals in females	183	177–198 (N=19) 187.21 ± 6.55	176–199 (N=5) 190 ± 8.86
Subcaudals in males	89–90 89.5 ± 0.71	83–107 (N=31) 94.65 ± 6.21	87–92 (N=7) 89.7 ± 1.8
Subcaudals in females	76	72–93 (N=16) 83.38 ± 6.11	77–94 (N=4) 85.25 ± 8.5
Preoculars	2–3 usually 2	1–3 usually 2	1–3 (N=17) usually 1
Postoculars	2	2–4 usually 2	2–3 (N=11) usually 2
Supralabials	8	8	7–9 (N=20) usually 8
Supralabials entering orbit	4–5	3–4–5, 4–5, 4–5–6 usually 4–5	3–4–5, 4–5, 4–5–6 usually 4–5
Infralabials	10	9–12 usually 10	8–12 (N=20) usually 10
Temporal formula	1+1, 1+2, 2+1 usually 1+2	1+1, 1+2, 1+3, 2+2 usually 1+2	1+2, 2+2 (N=10) usually 1+2
Dorsal body blotches	54–72 63.25 ± 7.89	30–47* 38.8	42–54 (N=5) 49.6 ± 5.37
Maximum SVL in females	523 mm	648 mm	630 mm (N=5)
Maximum SVL in males	445 mm	639 mm	443 mm (N=10)
Tail length/body length in females	0.30 (N=1)	0.21–0.26 (N=15) 0.24 ± 0.02	0.3–0.36 (N=4) 0.34 ± 0.03
Tail length/body length in males	0.36–0.39 (N=2) 0.37 ± 0.02	0.23–0.30 (N=28) 0.27 ± 0.02	0.3–0.48 (N=7) 0.37 ± 0.05

* Data from Duellman (1958) for specimens of “ Leptodeira septentrionalis ornata ” from western Ecuador .