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Chironius scurrulus (Wagler, 1824) Material examined. ECUADOR • 1 juvenile; Napo Province, WWS, Lodge; 00.6756°S, 077.6012°W; 1504 m a.s.l.; 11 March 2017; QCAZ 15407. Identification. This species has a color pattern of irregular, reddish-brown splotches on a blackish ground color. They have only 10 dorsal scale rows at midbody. Dorsal scales are smooth, the cloacal plate is undivided, and the ventral edges of the supralabial scales are light colored., Published as part of Camper, Jeffrey D., Torres-Carvajal, Omar, Ron, Santiago R., Nilsson, Jonas, Arteaga, Alejandro, Knowles, Travis W. & Arbogast, Brian S., 2021, Amphibians and reptiles of Wildsumaco Wildlife Sanctuary, Napo Province, Ecuador, pp. 729-751 in Check List 17 (3) on page 743, DOI: 10.15560/17.3.729

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). 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 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). 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 L. misinawui were encountered active at night near roots of shrubs, near trails, and in a small water pool. * Data from Duellman (1958) for specimens of “ Leptodeira septentrionalis ornata ” from western Ecuador.

1. Synophis niceforomariae Pyron, Arteaga, Echevarr��a, & Torres-Carvajal, 2016 Range. Andean highlands of north-central Colombia, Antioquia department, near Medell��n, ~ 1300���1700m, possibly with populations south of Medell��n, ~ 900m. Description. Adult size ~ 410mm SVL and ~ 210mm TL in males, smooth nuchal scales; no nuchal collar; intranasals in contact; 2 postoculars; 10���12 infralabials; 8 supralabials; 184���193 ventrals; 127���131 subcaudals; and 19-19-17 dorsal scale-rows with single, weak keel. Notes. Includes specimens referred to as S. bicolor and S. cf. bicolor by previous authors (Nic��foro - Mar��a 1970; Pyron et al. 2015; Torres-Carvajal et al. 2015)., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on page 302, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Niceforo - Maria, H. (1970) Contribucion al conocimiento de los ofidios de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 210, 1 - 6.","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058","Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533"]}

Synophis niceforomariae sp. nov. Holotype. MHUA 14577 (Fig. 2), adult male collected 24 November 2007 at Finca La Esperanza, Valle de La Manguita, Municipio Amalfi, Departamento Antioquia, Colombia (N6.978611, W-75.044444; 1394m). Paratypes. MHUA 14413 (Fig. 3), adult collected at Municipio Anori, Departamento Antioquia, Colombia (N6.987778, W-75.044444; 1320m); MLS 2072, adult male collected at Municipio Campamento, Departamento Antioquia, Colombia (N6.979498, W-75.296602; 1656m). Synophis calamitus BMNH 1940.2.30.31 - 8 2 162 118 21 19 17 408 241 M Synophis calamitus CAS 23612 11 8 2 166 100 19 19 17 186 80 F Synophis calamitus KU 164208 9 8 1 163 125 21 19 17 142 73 - Synophis calamitus KU 197107 9 7 1 166 110 21 19 17 149 74 F Synophis calamitus MCZ R-164530 11 9 2 164 116 - 19 17 367 208 - Synophis calamitus MZUTI 3694 11 9 2 166 118 23 19 17 462 265 M Synophis calamitus QCAZ 10453 10 8 2 160 - 19 19 16 502 - F ������continued on the next page Synophis lasallei EPN S.974 - - 2 156 116 - 21 - 175 90 M Synophis lasallei EPN S.975 - - 2 155 119 - 21 - 354 201 M Synophis lasallei FHGO 6489 11 8 2 147 111 23 21 21 153 86 M Synophis lasallei FMNH 81313 - - 2 154 112 - 21 - 292 158 F Synophis lasallei MCZ R-156873 11 7 1 147 115 - - - 412 206 - Synophis lasallei MLS / CJSP - - 2 144 101 - - - 300 170 M ������continued on the next page Species Collection IL SL PO V SC D1 D2 D3 SVL TL Sex Synophis lasallei MZUTI 4181 11 9 2 156 29 21 21 19 272 42 M Synophis lasallei USNM 233061 11 9 2 156 124 - 21 - 285 160 M Synophis lasallei USNM 233062 11 8 2 153 126 - 22 20 360 200 M Synophis lasallei USNM 233063 11 8 2 151 86 23 21 19 308 197 M Synophis lasallei USNM 233064 11 8 2 151 - - 21 19 270 150 - Etymology. Named after the French priest and naturalist Hermano Nic��foro Mar��a (born Antoine Rouhaire Siazaude), who established important natural-history collections at the Instituto de la Salle in Bogot��. While numerous taxa have been named mariarum, nicefori, or some derivative (see Adler 1989), he requested in a 1968 letter that he be addressed in print using his full name, Hermano Nic��foro Mar��a (Rodriguez 2002). Thus, it seems appropriate to use the specific epithet niceforomariae. Diagnosis. Synophis niceforomariae can be differentiated from all other similar or related species by the combination of an unmodified vertebral scale row with a single weak keel; smooth nuchal scales; absence of a nuchal collar in adults; presence of a loreal; intranasals in contact; fused prefrontals; rostral concave; anal single; no apical pits; 2 postoculars; 10���12 infralabials; 8 supralabials; 184���193 ventrals; 127���131 subcaudals; and 19-19- 17 dorsal scale-rows. Description. Small-sized, long-tailed snakes (~ 400mm SVL, 200mm TL) with slender bodies and head distinct from neck. Eye large (>1/3 head height), bulbous, and black in life, with pupil not easily distinguishable from iris. Pupil round in preservative (though this may be an effect of fixation). Dorsum coloration grayish-brown with iridescent sheen in life and preservation, no light-colored nuchal collar in adults, and posterior supralabials mostly unpigmented (Remarks. This species was referred to as Synophis cf. bicolor by Pyron et al. (2015). The cleared-and-stained skull of the male holotype was prepared by Martinez (2011), with illustrations of the cranium and mandible (her Figs. 15, 18, & 22), but the hemipenis was not illustrated or described in detail. Condition of the vertebrae, which are heavily modified in Emmochliophis and some Synophis (Fritts & Smith 1969; Savitzky 1974; Hillis 1990) unknown, pending skeletal preparation, micro-CT scanning, or X-rays. Has a very small known range to the east of Yarumal, Colombia, at ~ 1300���1600m around Campamento, Anori, and Amalfi. Two reportedly similar specimens in the Instituto de Ciencias Naturales de la Universidad Nacional de Colombia (ICN) were collected slightly to the south of the populations sampled here, at lower elevations (~ 900���1300m; Table 2). We did not examine these individuals, but it seems likely that they are also S. niceforomariae, and that the range of the species is slightly larger (Fig. 4). With this new species and the re-assignment of numerous specimens examined by Pyron et al. (2015) and Torres-Carvajal et al. (2015), the known species limits of many Synophis taxa have changed. In particular, Pyron et al. (2015) assigned specimens of S. bogerti, S. calamitus, and S. niceforomariae to S. bicolor and assigned specimens of S. zamora to S. bicolor and S. lasallei. Similarly, Torres-Carvajal assigned specimens of S. niceforomariae to S. bicolor. Using the updated information from the combined datasets (Tables 1, 2), we provide a checklist with updated summaries of geographic, elevational, and external morphological data useful for classifying the 9 species of Synophis known to date, Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on pages 295-302, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Society for the Study of Amphibians and Reptiles. & Adler, K. (1989) Contributions to the history of herpetology. S. S. A. R., Oxford, Ohio, 202 pp.","Rodriguez, G. (2002) El naturalista Antoine Rouhaire (Hermano Niceforo Maria) y el desarrollo de zoologia sistematica en Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 26, 229 - 238.","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058","Martinez, P. M. S. (2011) Comparative anatomy of the tribe Nothopsini (Serpentes, Dipsadidae), Thesis mestrado. Universidade de Sao Paulo, Sao Paulo, 159 pp.","Fritts, T. H. & Smith, H. N. (1969) A new genus and species of snake from western Ecuador. Transactions of the Kansas Academy of Science, 72, 60 - 66. http: // dx. doi. org / 10.2307 / 3627049","Savitzky, A. H. (1974) The relationships of the xenodontine colubrid snakes related to Ninia. MS thesis. University of Kansas, Lawrence, Kansas, 64 pp.","Hillis, D. M. (1990) A new species of xenodontine colubrid snake of the genus Synophis from Ecuador and the phylogeny of the genera Synophis and Emmochliophis. Occasional Papers of the Museum of Natural History, University of Kansas, 135, 1 - 9.","Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533"]}

6. Synophis insulomontanus Torres-Carvajal, Echevarr��a, Venegas, Ch��vez, & Camper, 2015 Range. Amazonian Andean slopes of central Peru, ~ 1100���1800m. Description. Adult size ~ 340���380mm SVL and ~ 180���190mm TL (assuming> 340mm SVL is adult, and CORBIDI 10418 has a truncated tail), keeled nuchal scales; no nuchal collar; intranasals in contact; 2 postoculars; 10���11 infralabials; 8���9 supralabials; 147���152 ventrals; 103���109 subcaudals; and (20-22)���(19-21)-19 dorsal scalerows, with single, heavy keel. Notes. Known only from the holotype and paratypes., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on pages 307-308, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533"]}

4. Synophis calamitus Hillis, 1990 Range. Pacific Andean slopes of central and northern Ecuador, primarily in Pichincha, Santo Domingo de los Ts��chilas, and Cotopaxi provinces, ~ 1000���2300m. Populations also occur in Carchi province (~ 1900m), and possibly Chimborazo province (~ 800m; see below). Description. Adult size ~ 330���510mm SVL and ~ 190���280mm TL (assuming> 330mm SVL is adult), smooth nuchal scales; no nuchal collar; intranasals usually in contact but occasionally divided by rostral; usually 2 postoculars but occasionally 0 or 1; 8���11 infralabials; 7���9 supralabials; 157���166 ventrals; 96���125 subcaudals including possibly truncated tails, 100���113 in females, 109���118 in males; and (19���23)-19-17 dorsal scale-rows with single, weak keel. Notes. Includes a much larger range of specimens identified as Synophis aff. bicolor from the western Andes by Pyron et al. (2015). The specimen from Carchi province in northern Ecuador (QCAZ 5847) is morphologically identical to other S. calamitus populations (Torres-Carvajal et al. 2015), but genetically distinct from the central Ecuadorean populations (Figs. 1, 4). There is thus intraspecific genetic diversity in S. calamitus, possibly related to the R��o Mira as a geographic barrier to gene flow. The southernmost specimen, from Chimborazo province (CAS 23612; Fig. 4), resembles both S. calamitus and S. zaheri. We refer it here to S. calamitus, for which it is a southern range extension, but it may represent a northern range extension for S. zaheri or a new species. Collecting effort in this locality should be a priority., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on page 307, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Hillis, D. M. (1990) A new species of xenodontine colubrid snake of the genus Synophis from Ecuador and the phylogeny of the genera Synophis and Emmochliophis. Occasional Papers of the Museum of Natural History, University of Kansas, 135, 1 - 9.","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058","Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533"]}

2. Synophis bicolor Peracca, 1896 Range. Chocoan lowlands of northwestern Ecuador, and presumably southwestern Colombia, ~ 200��� 300m. Description. Adult size ~ 360���410mm SVL and ~ 230���250mm TL in males, smooth nuchal scales; no nuchal collar; intranasals in contact; 2 postoculars; 9���11 infralabials; 8 supralabials; 174���183 ventrals; 129���143 subcaudals; and 19-19-17 dorsal scale-rows with single, weak keel. Synophis calamitus BMNH 1940.2.30.31 R��o Solaya, Ecuađor - 0.010213 -78.819510 1008 Synophis calamitus CAS 23612 Chimborazo, Naranjapata, Ecuađor -2.266667 -79.083333 763 Synophis calamitus KU 164208 9 km SE Tanđayapa, Pichincha Province, Ecuađor - 0.047404 -78.632804 2169 Synophis calamitus KU 197107 4 km SE Tanđayapa, Pichincha Province, Ecuađor - 0.012514 -78.650697 1889 Synophis calamitus MCZ R-164530 Ecuađor, Pichincha, Tanđapi -0.419803 -78.801132 1714 Synophis calamitus MZUTI 3694 Tambo Tanđa, Ecuađor - 0.020108 -78.651012 2048 Synophis calamitus QCAZ 10453 Cotopaxi: Naranjito, Bosque Integral Otonga -0.417820 -78.988030 1655 Synophis calamitus QCAZ 10453 Naranjito, Bosque Integral Otonga (BIO) -0.415967 -79.004800 2110 Synophis calamitus QCAZ 1136 Chiriboga -0.224530 -78.767750 1955 ������continued on the next page Species Collection Number Locality Latitude Longitude Elevation ������continued on the next page Notes. Populations representing S. bogerti, S. calamitus, S. niceforomariae, and S. zamora have previously been referred to as S. " bicolor " (e.g., Bogert 1964; Pyron et al. 2015; Torres-Carvajal et al. 2015)., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on pages 302-303, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Peracca, M. G. (1896) Nuovo genere di Colubride aglifo dell'America meridionale. Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universita di Torino, 11, 1 - 2.","Bogert, C. M. (1964) Snakes of the genera Diaphorolepis and Synophis and the colubrid subfamily Xenoderminae. Senckenbergiana Biologica, 45, 509 - 531.","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058","Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533"]}

7. Synophis zamora Torres-Carvajal, Echevarr��a, Venegas, Ch��vez, & Camper, 2015 Range. Amazonian Andean slopes of southeastern Ecuador, Zamora and Morona-Santiago provinces, ~ 800��� 1800m. Description. Adult size ~ 310���460mm SVL and ~ 150���210mm TL, keeled nuchal scales; no nuchal collar; intranasals in contact; 2 postoculars; 9���11 infralabials; 8���9 supralabials; 147���154 ventrals; 88���118 subcaudals; and (19���21)-(18���21)-(17���18) dorsal scale-rows with single, heavy keel. Notes. Includes several specimens referred to S. bicolor and S. lasallei by Pyron et al. (2015)., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on page 308, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058"]}

8. Synophis bogerti Torres-Carvajal, Echevarr��a, Venegas, Ch��vez, & Camper, 2015 Range. Amazonian Andean slopes of central Ecuador, ~ 1000���2000m. Description. Adult size ~ 370���410mm SVL and ~ 180���200mm TL in males and 420���540mm SVL and 220��� 240mm TL in females (assuming> 370mm SVL is adult), keeled nuchal scales; no nuchal collar; intranasals in contact; 2 postoculars; 10���11 infralabials; 8���9 supralabials; 157���168 ventrals; 103���117 subcaudals; and 19-(17��� 19)-17 dorsal scale-rows at midbody, with single, heavy keel. Notes. Includes specimens historically and recently referred to S. bicolor, S. aff. bicolor (eastern Andes), and S. lasallei by previous authors (Bogert 1964; Pyron et al. 2015)., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on page 308, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533","Bogert, C. M. (1964) Snakes of the genera Diaphorolepis and Synophis and the colubrid subfamily Xenoderminae. Senckenbergiana Biologica, 45, 509 - 531.","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058"]}

3. Synophis zaheri Pyron, Guayasamin, Pe��afiel, Bustamante, & Arteaga, 2015 Range. Pacific Andean slopes of southwestern Ecuador, El Oro province, ~ 800���900m. Description. Adult size ~ 350���370mm SVL and ~ 180���190mm TL in males, smooth nuchal scales; no nuchal collar; intranasals in contact; 2 postoculars; 8���9 infralabials; 8 supralabials; 166���169 ventrals; 111���112 subcaudals; and 19-19-17 dorsal scale-rows with single, weak keel. Notes. Known only from the holotype and paratype., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on page 307, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058"]}

9. Synophis plectovertebralis Sheil & Grant, 2001 Range. Pacific Andean slopes of Valle del Cauca, Colombia, ~ 1800m. Description. Adult size ~ 200���210mm SVL and ~ 80���100mm TL, smooth nuchal scales; nuchal collar; intranasals in contact; 1 postocular; 7���8 infralabials; 7���8 supralabials; 144���147 ventrals; 79���91 subcaudals; and 19-19-19 dorsal scale-rows at midbody, with single, weak keel. Notes. Known only from the holotype and paratype. Not included in our phylogenetic analyses (Fig. 1); its placement is thus unknown. With this re-assessment of species boundaries and taxonomic reassignment of nearly all known specimens of Synophis (Figs. 1���4, Tables 1,2), the morphological limits and geographic ranges of the known, extant species are tentatively stabilized. There are likely additional cryptic species in some taxa (e.g., S. lasallei), and there are almost certainly additional new species to be discovered. Our results here should clarify lingering confusion over most species limits, and provide a robust basis for future work. The known species can be informally grouped into at least three species groups. The first is the S. bicolor group (S. bicolor, S. niceforomariae), distributed in the northern Andes, with higher ventral and subcaudal counts (174���193 and 127���143), weak keels, more slender bodies and lighter colorations (Figs. 2, 3; see Fig. 4 in Pyron et al. 2015). The second is the S. calamitus group (S. zaheri, S. calamitus), distributed on the Pacific Andean slopes of Ecuador, with intermediate counts (157���169 and 96���125), stronger keels, more robust bodies, and both lighter (S. zaheri) and darker (S. calamitus) colorations (see Figs. 3, 8, & 9 from Pyron et al. 2015 and Fig. 5 from Torres- Carvajal et al. 2015). The third is the S. lasallei group (S. lasallei, S. insulomontanus, S. zamora, and S. bogerti), with lower to intermediate counts (144���168 and 88���126), heavy keels, and darker colorations (see Figs. 3 & 9 from Pyron et al. 2015 and Fig. 5 from Torres-Carvajal et al. 2015). The placement of S. plectovertebralis is uncertain, and we suggest it may form a fourth group with small body sizes (~ 200mm SVL), low counts (144���147 and 79��� 91), nuchal collars, and weak keels. The phylogenetic placement of this group is not known, and it could potentially include Emmochliophis (E. miops was originally described in Synophis), which also have low counts and nuchal collars, but lack a loreal. The first three groups are also at least partially distinguishable by hemipenial morphology. In all known diaphorolepidine species, the hemipenis is bilobed, semicalyculate, and semicapitate, relatively stout and bulbous, and covered in large spines or hooks near the base. The organ is not markedly different between Diaphorolepis wagneri, Synophis bogerti and S. lasallei (Bogert 1964; Zaher 1999), and thus does not appear to strongly differentiate individual species, though no quantitative analysis sampling multiple individuals per species has been performed. The organ has also not been described in detail or illustrated for any species in the S. bicolor or S. plectovertebralis groups. In the S. calamitus group (S. calamitus QCAZ452; S. zaheri MZUTI3353, 3355), the lobes are stout and blunt, approximately one-half the length of the hemipenial body, which is ornamented with numerous spines, none of which are noticeably enlarged. In contrast, the S. lasallei group is diagnosed by lobes which are more slender than the hemipenial body, and approximately equal to its length. All species have a significantly enlarged spine near the base of the hemipenial body, either on the right side in sulcate view (S. lasallei USN233062; S. insulomontanus CORBIDI13940) or the left side (S. zamora QCAZ9174; S. bogerti QCAZ12791). An extensive comparison of more hemipenial material may reveal more significant differences that are useful for diagnosing species. Similarly, the vertebral condition of many of the newer species is not known. Diaphorolepidine species are generally characterized by heavy modification of the pre- and post-zygapophyses (Bogert 1964; Fritts & Smith 1969; Savitzky 1974; Hillis 1990; Sheil & Grant 2001). Examining this condition requires either destructive dissection, or expensive microCT scanning. A comparative analysis of material using the latter technique will likely prove invaluable for diagnosing species and determining species boundaries, as well as perhaps understanding the function of this unique adaptation. Finally, snakes in the genus Synophis have recently been referred to as "fishing snakes" by various sources, such as the Reptile Database (http://www.reptile-database.org/), the IUCN Red List (http://www.iucnredlist.org/), Torres-Carvajal et al. (2015), and various news outlets covering the discoveries of Pyron et al. (2015) and Torres- Carvajal et al. (2015). However, the origin of this name is unclear, as no known species are ichthyophagous. The name appears to originate from The Elsevier Dictionary of Reptiles (Wrobel 2004), but no etymology is given. The only known diet items are gymnophthalmid lizards in Emmochliophis miops and Synophis plectovertebralis (Sheil 1998; Sheil & Grant 2001). Thus, we suggest the Standard English Name Andean Shadow Snakes and Standard Spanish Name Culebras Andinas de la Sombra for the genus Synophis, based on their dark color, nocturnal habits, and secretive nature. For the species, the Standard English Names would be Nic��foro Mar��a's Shadow Snakes (S. niceforomariae), Bicolored Shadow Snakes (S. bicolor), Zaher's Shadow Snakes (S. zaheri), Calamitous Shadow Snakes (S. calamitus), La Salle's Shadow Snakes (S. lasallei), Mountain Shadow Snakes (S. insulomontanus), Zamoran Shadow Snakes (S. zamora), Bogert's Shadow Snakes (S. bogerti), and Braided Shadow Snakes (S. plectovertebralis). Suggested Standard Names in Spanish are, respectively, Culebras Andinas de la Sombra de Nic��foro Mar��a, Culebras Andinas de la Sombra bicolores, Culebras Andinas de la Sombra de Zaher, Culebras Andinas de la Sombra calamitosas, Culebras Andinas de la Sombra de La Salle, Culebras Andinas de la Sombra monteses, Culebras Andinas de la Sombra de Zamora, Culebras Andinas de la Sombra de Bogert, and Culebras Andinas de la Sombra trenzadas., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on pages 308-309, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Sheil, C. A. & Grant, T. (2001) A new species of colubrid snake (Synophis) from western Colombia. Journal of Herpetology, 35, 204 - 209. http: // dx. doi. org / 10.2307 / 1566109","Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058","Torres-Carvajal, O., Echevarria, L. Y., Venegas, P. J., Chavez, G. & Camper, J. D. (2015) Description and phylogeny of three new species of Synophis (Colubridae, Dipsadinae) from the tropical Andes in Ecuador and Peru. Zookeys, 153 - 179. http: // dx. doi. org / 10.3897 / zookeys. 546.6533","Bogert, C. M. (1964) Snakes of the genera Diaphorolepis and Synophis and the colubrid subfamily Xenoderminae. Senckenbergiana Biologica, 45, 509 - 531.","Zaher, H. (1999) Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History, 1 - 168.","Fritts, T. H. & Smith, H. N. (1969) A new genus and species of snake from western Ecuador. Transactions of the Kansas Academy of Science, 72, 60 - 66. http: // dx. doi. org / 10.2307 / 3627049","Savitzky, A. H. (1974) The relationships of the xenodontine colubrid snakes related to Ninia. MS thesis. University of Kansas, Lawrence, Kansas, 64 pp.","Hillis, D. M. (1990) A new species of xenodontine colubrid snake of the genus Synophis from Ecuador and the phylogeny of the genera Synophis and Emmochliophis. Occasional Papers of the Museum of Natural History, University of Kansas, 135, 1 - 9.","Wrobel, M. (2004) Elsevier's dictionary of reptiles: in Latin, English, German, French and Italian. Elsevier, Amsterdam; Oxford, 758 pp.","Sheil, C. A. (1998) Emmochliophis miops: Redescription of Synophis miops (Boulenger, 1898). Journal of Herpetology, 32, 604 - 607. http: // dx. doi. org / 10.2307 / 1565222"]}

5. Synophis lasallei (Nic��foro-Mar��a, 1950) Range. Amazonian Andean slopes, from central Colombia to central Ecuador, ~ 500���1500m. The type locality ("N of Alb��n, cen. Cundinamarca Dept., cen. Colombia ") is difficult to georeference accurately for precise location and elevation. Description. Adult size ~ 270���410mm SVL and ~ 150���210mm TL (assuming> 270mm SVL is adult and MZUTI 4181 and USNM 233063 have truncated tails), keeled nuchal scales; no nuchal collar; intranasals in contact; usually 2 postoculars, but occasionally 1; 11 infralabials; 7���9 supralabials; 144���156 ventrals; 101���126 subcaudals omitting truncated tails, 111���126 not including holotype MLS /CJP with 101, which may also be truncated but was not examined; and (21���23)-(21���22)-(19���20) dorsal scale-rows with single, heavy keel. Notes. Some specimens of S. bogerti and S. zamora were referred to S. lasallei by Pyron et al. (2015), prior to the description of those taxa. The only specimen of S. lasallei sequenced here is from the southernmost known population in Ecuador. It is possible that additional cryptic species are present in this taxon, such as the Colombian or northern Ecuadorean populations., Published as part of Pyron, R. Alexander, Arteaga, Alejandro, Echevarr��a, Lourdes Y. & Torres-Carvajal, Omar, 2016, A revision and key for the tribe Diaphorolepidini (Serpentes: Dipsadidae) and checklist for the genus Synophis, pp. 293-320 in Zootaxa 4171 (2) on page 307, DOI: 10.11646/zootaxa.4171.2.4, http://zenodo.org/record/158885, {"references":["Pyron, R. A., Guayasamin, J. M., Penafiel, N., Bustamante, L. & Arteaga, A. (2015) Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of Synophis from the Pacific Andean slopes of southwestern Ecuador. Zookeys, 109 - 147. http: // dx. doi. org / 10.3897 / zookeys. 541.6058"]}