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Sceloporus huichol sp. nov. Sceloporus torquatus melanogaster Zweifel. 1959. Amer. Mus. Novit. (1953):3–4. Sceloporus sp. Martínez-Méndez & Méndez-de la Cruz. 2007. Zootaxa (1609):53–68. urn:lsid:zoobank.org:pub: 4E3A7D0C-BB24-4407-9304-F90B589B2F6E Holotype. MZFC �20633 (Fig. 1), adult ♀ from municipality of Bolaños, highway Bolaños-Tuxpan de Bolaños, 2485 m, 21.91774 N, - 103.87735 W, Jalisco, Mexico (Fig. 2). Collected in pine-oak forest by P. Ponce Campos, 13 June 2003, original field number JAC 23414. Paratypes (20). Mexico: Jalisco: Municipality of Bolaños, highway Bolaños-Tuxpan de Bolaños, 2400 m, 21.89802 N, - 103.86037 W, 11 June, 2003 (UTA R-55432, ♀); same data as holotype (MZFC �20634–37, UTA R-55435–36; three ♀, and one ♂; and one ♂, and one ♀, respectively); Jalisco: Municipality of Bolaños, cell phone antenna, 2549 m, 21.89285 N, - 103.86502 W, 13 June, 2003 (UTA R-55433–34, ♀ and ♂, respectively); 7 km N, 6 km W Bolaños, 2345 m, 21.886972 N, - 103.84033 W (ENCB 14315–16, ♀ and ♂, respectively); Nayarit: Sierra Alica, highway Huajimic–Tepic, 1859 m, 21.67361 N, - 104.40736 W, 14 June, 2003 (UTA R-55437, ♀); Nayarit: 9 km SE Huajimic, 21.64458 N, - 104.276374 W (CNAR5819–1, 5819–2, 5819–5, 5819–7, 5819–8, 5819–9, 5819– 12, 5819–13; four ♂ and four ♀). All localities are located in pine-oak forest. Diagnosis. A species of the torquatus group sensu stricto (i.e., clade A of Martínez-Méndez & Méndez-de la Cruz 2007; Wiens et al. 2010), more closely related to S. melanogaster than to any other described species in the group (Martínez-Méndez & Méndez-de la Cruz 2007). This species is characterized by the combination of the following characters: four to five supraocular scales in a single row; large dorsal scales, arranged in 28–30 rows from occiput to base of tail; femoral pores 14–21; ventral scales 42–51; scales around midbody 37–43; maximum snoutvent length (SVL) 105 mm; complete dark nuchal collar, 2.5–6 scales wide, with light paravertebral bands reaching behind the parietal region; white marks on head scales; limbs barred. The new species differs from S. melanogaster by having paravertebral light bands running from nuchal collar reaching behind the parietal region (not forming continuous paravertebral light bands from nuchal collar to the parietal region in S. melanogaster); having white marks on center or edges of head scales and postoccipital region (lacking in S. melanogaster); having 37–43 scales at midbody (S. melanogaster with 31–46 scale rows); maximum SVL for both sexes is 105 mm (S. melanogaster larger, reaching 131 mm SVL in ♂ [MZFC �28087] and 120 mm in ♀ [CNAR �6090]); ♀ S. melanogaster S. torquatus by having always complete nuchal collar, 4–6 scales wide (usually complete nuchal collar, 4–5 scales wide in S. torquatus); usually having white marks on top of head (rarely present in S. torquatus); by having always barred limbs (limbs rarely barred in S. torquatus); maximum SVL 105 mm (S. torquatus larger, ♂ over 111 mm SVL [MZFC 3954] and ♀ over 105 mm SVL [MZFC 3960]). Differing from S. binocularis by having white marks on top of head (lacking in S. binocularis); by having complete dark nuchal collar (incomplete in S. binocularis); possessing more femoral pores 14–21 (12–16 in S. binocularis); more scales around midbody 37–43 (29–36 in S. binocularis); smaller size, maximum SVL for S. huichol sp. nov. 105 mm (120 mm for S. binocularis MZFC 7421). Differs from S. mikeprestoni by having a dark nuchal collar wide of 4–6 scales (3–4 in S. mikeprestoni); by having white marks on top of head (rarely present in S. mikeprestoni); having barred limbs (not barred in S. mikeprestoni). Differs from S. madrensis by having white marks on top of head (rarely present in S. madrensis); having barred limbs (not barred in S. madrensis); by having fewer ventrals 42–51 (44–56 in S. madrensis); more scales around midbody 37–43 (34–41 in S. madrensis) and having more femoral pores 14–21 (13–18 in S. madrensis). Differs from S. bulleri by having white markings on top of head (lacking in S. bulleri); having barred limbs (not barred in S. bulleri); fewer dorsal scales, 28–30 (35–44 in S. bulleri); fewer ventral scales, 40–46 (47–61 in S. bulleri) and by being smaller, SVL 105 mm (116 mm in S. bulleri). Differing from S. insignis by having a wider nuchal collar, 4–6 scales wide (2–3 in S. insignis); fewer dorsal scales, 28–30 (37–47 in S. insignis); fewer ventral scales, 42–51 (54–61 in S. insignis); fewer scales at midbody, 37–43 (40–49 in S. insignis) and being larger, 105 mm SVL (99 mm in S. insignis). Differing from S. jarrovii being larger SVL, 105 mm (89 mm in S. jarrovii); fewer dorsals scales, 28–30 (38–45 in S. jarrovii); having 37–43 scales around midbody (43–58 in S. jarrovii); fewer ventral scales 42–51 (54–64 in S. jarrovii); a wider dark nuchal collar 4–6 scales (2.5– 4 in S. jarrovii); having barred arms and legs (only barred arms in S. jarrovii). Sceloporus huichol sp. nov. differs from all other species in the poinsettii group (i.e., clade B in Martínez-Méndez & Méndez-de la Cruz 2007; Wiens et al. 2010) by having enlarged supraocular scales in a single series, while all others, except S. serrifer Cope and S. macdougalli Smith & Bumzahem, have two series. Sceloporus huichol sp. nov. differs from S. serrifer by having a shorter SVL, 105 mm (112 mm in S. serrifer); having more scales at midbody, 37–43 (29–35 in S. serrifer); having more ventral scales, 42–51 (36–47 in S. serrifer); having more femoral pores, 14–21 (S. serrifer with 8–14); having a wider nuchal collar, 4–6 scales wide (2–4 in S. serrifer); and by having barred limbs (not present in S. serrifer). Sceloporus huichol sp. nov. differs from S. macdougalli by having more ventral scales, 42–51 (34–43 in S. macdougalli); more scales at midbody, 37–43 (31–35 in S. macdougalli); more femoral pores, 14–21 (12–17 in S. macdougalli); by having barred limbs (only arms are barred in S. macdougalli); wider nuchal collar, 4–6 scales wide (narrower, 3–4 scales wide in S. macdougalli); and by having paravertebral bands reaching parietal region (not present in S. macdougalli). Description of holotype. Dorsal head scales smooth, not pitted except for rostral, postrostrals, and internasals; frontal divided, posterior half contacting interparietal, anterior half contacting two prefrontals; prefrontals pentagonal, in contact medially, not separated by frontonasal; three frontonasals; lateral frontonasals broadly contacting median frontonasal, median frontonasal hexagonal, wider than high, lateral frontonasals hexagonal but higher than wide; five total internasals, 3 on right, 2 on left side; 6 scales contacting rostral between anterior supralabials; nasals separated from postrostrals by one scale on both sides; two canthals on each side, anterior smaller; 6–6 superciliaries, all separated from supraoculars by one row of small scales; enlarged supraoculars 4–4; one parietal and one frontoparietal on each side, frontoparietals not in contact medially; scales on side of head pitted, except for some lorilabials and labials; one large subnasal on both sides; one loreal on each side; one preocular larger than loreal on each side; one subocular on each side of head; lorilabials in 2 rows; supralabials 5–5, 4th below middle eye (Fig. 3). Outer row of labiomentals separated from mental; first infralabial contacting mental and first postmental; 6–6 infralabials. Dorsal scales mucronate; median keel stronger on dorsolateral scales; variable serrations on each side of scale point; dorsals 29; ventrals 46, smooth; scales around midbody 40. Femoral pores 17–17, separated medially by seven ventral scale rows; lamellae on fourth toe 20–20. Measurements. SVL 89.3 mm; tail 70.0 mm (regenerated); tibia 20.0 mm; snout to posterior edge of ear 22.3 mm; width of head at level of anterior edge of ear 19.7 mm. Color in preservative. Dorsum of head black, fading towards snout, with cream lines beneath eyes; interparietal black with pale cream dot on center of scale and light edges; five cream marks radiating from interparietal, one on posterior frontal and posterior part of anterior frontal, two extending to lateral side covering parietals, and two radiating upward from posterior corners of base of interparietal (not in center of scales as in S. serrifer); small pale spot on two scales immediately behind interparietal; prefrontals, frontonasals, internasals, and scales in nasal region brown; each supraocular scale dark with small pale dot in center, which is most evident on last posterior supraocular; subnasals, canthals, preoculars and scales below cream, extending to lip, forming line below eye and extending across the tympanum to neck and nuchal collar (Fig. 1); interrupted black band from behind interparietal to nuchal collar forming pattern on neck consisting of a black line (covering about two adjacent half scales) on middorsum of neck and two pale olive-green lateral bands about two scales wide. Nuchal collar black, extending from shoulders to anterior third of dorsum, forming “V”, from two to three and one-third scales wide at widest part on side of neck; posterior margin of dark nuchal collar bordered by cream band about one scale wide from shoulder to side of dorsum, not completely enclosing black nuchal collar; nuchal collar extends 15 scales on dorsum; dorsum almost uniformly olive-green posterior to nuchal collar, first row of scales behind collar slightly paler than other dorsals; fore and hind limbs barred (brown bands on olive green background), ventral surfaces of limbs not barred; venter of head cream with narrow transverse grey bands becoming poorly defined posteriorly; venter of body mostly immaculate cream, including base of tail, except for sides of venter with pale blue belly patches running parallel to body between fore and hind limbs. Color in life (based on paratype UTA R-55437, Fig. 4). Dorsum of head dark brown with cream line beneath eye; interparietal dark brown with pale cream dot from center to posterior end of scale and continuous with pale spot on first two scales behind interparietal (holotype similar); each supraocular scale dark with small pale dot on center of scale; canthal and subocular regions cream, projecting behind head forming “V” shaped mark on sides of head, region behind eye dark brown; black band from behind interparietal to nuchal collar forming pattern on neck consisting of black line (includes vertebral scale row and adjacent half scales on either side) on middle of neck and two pale olive green paravertebral bands about two scales wide. Nuchal collar black, extending from shoulders to anterior third of dorsum, forming “V”, from two to three and one-third of scale wide at widest part; posterior margin of nuchal collar bordered by cream band, one scale wide from shoulder to side of dorsum, not completely enclosing black nuchal collar; nuchal collar not present on venter; dorsal coloration pale brown on center of dorsum, grey on both sides, some dorsal scales with black markings on posterior tip forming irregular, discontinuous black cross bands (five from behind nuchal collar to base of tail); front and hind limbs barred (black bands on grey background), ventral parts of limbs not barred; tail complete, banded with black and cream bands on grey background, background fading towards end of tail with only alternating black and cream bands remaining; venter of head pale grey with cream scattered spots on anterior part; pale bluish mark with cream spots on posterior part reaching anterior part of chest; venter cream from chest to base of tail, sides of venter with patch running parallel to body between front and hind limbs on each side of pale campanula (Smithe, 1975 color No.71). Juvenile coloration (coloration based on paratype UTA R-55432, Fig. 5, color in life). Dorsum of head brown with white dots on center of interparietal, parietals, frontoparietals, supraoculars, anterior frontal, lateral frontonasals and first dorsal scale; posterior frontal covered by white; a white line from canthal crossing eye to temporal region of head, black line running paralleled and above white line; rostral, upper labials, lorilabials, preocular, subocular forming white band on side of head and extending posteriorly to anterior edge of ear opening; five white marks on dorsum of neck, two on sides behind interparietal covering two scales bordered by tiny black lines, one on center of neck one scale wide, and two lateral in front insertion of arms on shoulder similar to others; few scattered white marks on sides of neck not larger than two scales; dorsum of body pale brown, dark nuchal collar bordered anteriorly and posteriorly by white, incomplete at middorsum, nuchal collar no more than four scales wide; dorsum of body with white and black dots scattered from behind nuchal collar to base of tail; fore and hind limbs barred with brown and white bands; tail barred similar to limbs; ventral surfaces of head and body pale bluish, venter of limbs and tail cream. Variation. The variation reported is based on 27 specimens (21 of these are the type and paratypes), of which 13 are adult ♀, eight adult ♂ and six juveniles; three juveniles are ♂ (sex verified by the presence of two large postcloacal scales); the six additional specimens that we examined are all adults. Dorsal anterior head scales pitted, eight specimens with pitted anterior frontal and supraocular scales (CNAR 5819–5, 5819–6, 5819–8, 5819–10, 5819–12, 5819–14, UTA R-55433 and 55437); prefrontals in contact except for nine specimens (ENCB 14315, 14316, CNAR 5819–2, 5819–5, 5819–7, 5819–8, 5819–9, 5819–12, UTA R-55437), six of these with azygous scale separating prefrontals (UTA �R-55437, CNAR 5819–2, 5819–5, 5819–9, ENCB �14315, 14316); one specimen with right frontonasal separated from the central by an azygous scale (UTA R-55437); internasals vary from two to five, arranged one on each side (CNAR 5819–14), two on each side (UTA R-55433, MZFC 20634, CNAR 5819–2, 5819–3, 5819–4, 5819–5, 5819–6, 5819–7, 5819–9, 5819–10), and the remaining irregularly arranged; nasal usually separated from rostral by one or two scales; superciliaries 6–6, only two specimens with 6–5 (UTA R-55437, CNAR 5819–4); one preocular on both sides (CNAR 5819–1, 5819–4, 5819–9, 5819–10, 5819–11, 5819–13, ENCB 14315, 14316), two preoculars on both sides (CNAR 5819–5, CNAR 5819–6, UTA �R-55437), and the remaining irregularly arranged; lorilabials usually in two rows, except for UTA R-55433 with 2–3 rows; supralabials 5–5 on each side, one specimen (CNAR 5819–14) 4–5, three specimens are damaged (CNAR 5819–4, 5819–7, 5819–9); infralabials 6–6 on each side, 6–5 (CNAR 5819–10), 6–7 (CNAR 5819–6), 5–6 (CNAR 5819−8), 7−7(CNAR 5819–1); lamellae under fourth toe 17–21; scale counts vary from: dorsals (28–30); scales around midbody 37–43; ventrals 42–51; femoral pores 14–21; scales between femoral pores 4–8. Measurements. SVL of 20 specimens 61.8–105.4 mm (larger specimen CNAR 5819–4); tail length 94.1–128.0 mm in seven paratypes having complete tail (CNAR 5819–1, 5819–5, 5819–6, 5819–7, 5819–11, MZFC 20634, UTA R-55437); head length 15.0–26.4 mm; head width 14.0–22.2 mm; tibia length 12.8–21.0 mm; SVL in juveniles 31.6–39.6 mm; tail broken in all juveniles; head length 10.5–12.0 mm; head width 7.8–9.5 mm; and tibia length 7.2–9.5 mm. Distribution. Sceloporus huichol sp. nov. inhabits the southern portion of the Sierra Madre Occidental, from San Juan volcano and Sierra de Álica in Nayarit, eastward to Sierra Los Huicholes in northern Jalisco. It occurs in pine-oak forest and is most often found on rocky outcropping and walls between 1859 and 2549 m above sea level. Etymology. The specific epithet is a patronym in honor of the Wixárika ethnicity, commonly referred to as Huicholes, a native people who have lived in the general area of this new species. Sometimes referred to as the last peyote guardians, who struggle to keep their traditions alive., Published as part of Flores-Villela, Oscar, Smith, Eric N., Campillo-García, Gustavo, Martínez-Méndez, Norberto & Campbell, Jonathan A., 2022, A new species of Sceloporus of the torquatus group (Reptilia: Phrynosomatidae) from West Mexico, pp. 286-296 in Zootaxa 5134 (2) on pages 288-292, DOI: 10.11646/zootaxa.5134.2.7, http://zenodo.org/record/6534998, {"references":["Martinez-Mendez, N. & Mendez-de la Cruz, F. R. (2007) Molecular phylogeny of the Sceloporus torquatus species-group (Squamata: Phrynosomatidae). Zootaxa, 1609, 53 - 68. https: // doi. org / 10.11646 / zootaxa. 1609.1.2","Wiens, J. J., Kuczynski, C. A., Arif, S. & Reeder, T. W. (2010) Phylogenetic relationships of phrynosomatid lizards based on nuclear and mitochondrial data, and a revised phylogeny for Sceloporus. Molecular Phylogenetics and Evolution, 54, 150 - 161. https: // doi. org / 10.1016 / j. ympev. 2009.09.008","Smithe, F. B. (1975) Naturalist's color guide. Part I. The American Museum of Natural History, New York, 18 pp."]}

Genus Salvadora Baird & Girard, 1853 Salvadora Baird & Girard, 1853: 104–105. Phimothyra Cope, 1861: 566. Type species Salvadora grahamiae Baird & Girard, 1853. Diagnosis Usually nine-plate supracephalic scaling on the head; enlarged rostral scale that intercedes between the internasals and sometimes has free edges; two nasal scales; preoculars and postoculars present; loreal scale simple or divided; smooth body scales sometimes with two apical pits; 17 rows of scales around the body on the anterior third of body and midbody; cloacal scale divided; subcaudal scales in two series; elongated head; large eyes; round pupil; long tail. Hemipenis simple with spines at the base, replaced midway by calyces that cover half the length of each organ: sulcus simple with a long basal spine. Normally from 9 to 15 maxillary teeth followed by a diastema and three enlarged teeth (9–15+ 3). Color pattern consisting of two longitudinal, dorsolateral and lateral lines on each side of the body and a pale vertebral line generally present. Etymology Derived from the Latin ‘ salvator ’, which means ‘savior’, and ‘ adora ’, which means ‘honor’, an explicit homage to the collector of the type species S. grahamiae, Col. J.D. Graham. Distribution Southern United States from southern California, western Nevada, Utah, to Arizona, New Mexico, to central Texas; in Mexico, the Baja California Peninsula and Sonora south to the lowlands of Chiapas and Guatemala excluding the Yucatán Peninsula. Remarks The phylogenetic hypothesis and the character revision reveal that there are two groups within the genus, as previously suggested by Smith (1938, 1941): mexicana and grahamiae., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on page 90, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Baird S. F. & Girard C. 1853. Catalogue of North American Reptiles in the Museum of the Smithsonian Institution. Part 1 Serpents. Smithsonian Institution, Washington D. C. https: // doi. org / 10.5962 / bhl. title. 5513","Cope E. D. 1861. Catalogue of the Colubridae in the Museum of the Academy of Natural Sciences of Philadelphia. Part 3. Proceedings of the Academy of Natural Sciences of Philadelphia 12: 553 - 566.","Smith H. M. 1938. Notes of the snakes of the genus Salvadora. The University of Kansas Science Bulletin 25: 229 - 237. https: // doi. org / 10.5962 / bhl. part. 1701","Smith H. M. 1941. Further notes on Mexican snakes of the genus Salvadora. Smithsonian Miscellaneous Collections 99: 1 - 12. Available from https: // repository. si. edu / handle / 10088 / 23709 [accessed 23 Jul. 2021]."]}

Salvadora intermedia Hartweg, 1940 Fig. 10 Salvadora intermedia Hartweg, 1940: 256–259. Salvadora intermedia richardi Smith, 1941: 6–9. Salvadora intermedia intermedia – Smith, 1941: 7. Diagnosis Rostral scale elongated; generally 8 supralabials (occasionally 9), the fourth and fifth normally in contact with the eye; 10–11 infralabials; preocular scale divided; a single loreal scale; prenasal scale in contact with or separated from the second supralabial; chinshields in contact or separated by two small scales; ventral scales 172 to 190; subcaudal scales 84 to 114; tail is 23 to 30% of the total length; 11 +3 maxillary teeth. Color pattern formed by a vertebral line three scales rows wide, yellowish in color and in contact with the parietal region; a pair of lateral lines is fused with the dorsolateral lines on first third of body, separated at second third of the body to tail; dorsolateral lines wide and dark, diverging at the neck, passing through the temporal region to eye. S alvadora intermedia differs from its congeners by having a divided preocular scale; fewer than 116 subcaudal scales; 8 supralabials, the fourth and fifth in contact with the eye; the second pair of chinshields in contact with each other or separated by a row of scales; and two dark dorsolateral lines that diverge at the neck, passing through the temporal to eye. Type material examined MEXICO • ♂, holotype of Salvadora intermedia intermedia; Guerrero, vicinity of Chilpancingo; W.W. Brown leg.; UMMZ 85733. MEXICO • ♂, holotype of Salvadora intermedia richardi; Puebla, 1 mile N of Tehuacán; 8 Aug. 1940; R.C. Taylor leg.; Collection E.H. Taylor and H.M. Smith No. 23470; UIMNH 25071. Distribution and conservation Endemic to Mexico. Distributed in central-southern Mexico from the vicinity of Chilpancingo, Guerrero, eastward through central Oaxaca and northward to southwestern Puebla (Fig. 10). Within its distribution it is found in at least two protected natural areas. Mexican legislation has listed this species as subject to Special Protection and the IUCN has it listed as a species of Least Concern. Remarks Salvadore intermedia richardi was described by Smith (1941), who based his description on a single specimen collected one mile from Tehuacán, Puebla (UIMNH 25071). Smith (1941) thought this specimen represented a disjunct population of the populations of S. intermedia at Guerrero. Smith (1941) defined S. intermedia richardi as having the prenasal scale separated from the second supralabial and by its number of ventral scales. However, we examined 37 individuals of S. intermedia along what appears to be a continuous distribution from Guerrero and through Oaxaca to southern Puebla, and found that only seven of these specimens had the condition of the prenasal scale separated from the second supralabial described by Smith (1941): three from Oaxaca (MZFC 04059, MZFC 23857, MZFC 23871), three from Guerrero (MCZ 33642, MZFC 02878, FMNH 109866), one from Puebla (MZFC 14384). Additionally, the number of ventral scales of these specimens falls within the range previously described for S. intermedia. Based on these findings, we conclude that these characters are part of the intrapopulational variation of S. intermedia and that S. i. richardi should be considered a junior synonym of S. intermedia as previously suggested by Davis & Dixon (1957)., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 103-104, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Hartweg N. 1940. Description of Salvadora intermedia, new species, with remarks on the grahamiae group. Copeia 1940 (4): 256 - 259. https: // doi. org / 10.2307 / 1438583","Smith H. M. 1941. Further notes on Mexican snakes of the genus Salvadora. Smithsonian Miscellaneous Collections 99: 1 - 12. Available from https: // repository. si. edu / handle / 10088 / 23709 [accessed 23 Jul. 2021].","Davis W. B. & Dixon J. R. 1957. Notes on Mexican snakes (Ophidia). The Southwestern Naturalist 2: 19 - 27. https: // doi. org / 10.2307 / 3669560"]}

Salvadora bairdi Jan in Jan & Sordelli 1860 Fig. 5 Salvadora bairdi Jan in Jan & Sordelli, 1860: pl. 3 fig. 2. Salvadora bogerti Smith, 1941: 2–6. Diagnosis Rostral without free edges, slightly elongated towards the back interceding between internasals; 8 supralabials, generally fourth and fifth touching the lower edge of the eye; 10 infralabials; preocular divided; a single loreal; prenasal may or may not be in contact with the second supralabial; a second pair of chinshields in contact with each other or sometimes separated by a row of scales;175–209 ventrals; 82–108 subcaudals; tail length 22 to 29% of the total length; maxillary teeth normally 9+3. Color pattern consists of a pale vertebral line three scales wide that begins on the neck and tapers to one scale wide on the last third of body; two pairs of dark longitudinal lines, dorsolateral lines two to four scales wide not touching the temporal region and connecting on the anterior part, forming a patch that separates them from the parietal region; lateral lines involving the third row of dorsal scales and may or may not be interrupted by an irregular pattern of patches in the posterior region of head. This species differs from its congeners by having a single loreal, edges of rostral scale not free, maxillary teeth 9 +3, and a pale vertebral line that extends to the neck without touching the temporal region. Etymology Named in honor of Spencer Fullerton Baird. Type material MEXICO • holotype of Salvadora bogerti; “Tehuantepec”, locality uncertain; USNM 30296. MEXICO • holotype of Salvadora bairdi; “Méxique” Guanajuato, restricted to Acámbaro, see Smith & Taylor (1950); Museo Civico di Storia Naturale di Milano, Westphal coll.; MSNM (destroyed). Distribution and conservation Endemic to Mexico; distributed from Sonora and Chihuahua southward through the states of Sinaloa, Durango, Zacatecas, Nayarit, Jalisco; in central Mexico through the Trans-Mexican Volcanic Belt in the states of Querétaro, Guanajuato, Michoacán, Guerrero, Estado de México, Hidalgo, Morelos, Puebla, and Veracruz (Fig. 5). Within its distribution, it is found in at least four protected natural areas. Mexican legislation has listed this species as subject to Special Protection and the IUCN has it listed as a species of Least Concern. Remarks The holotype of S. bogerti is a juvenile purportedly collected by F. Sumichrast in the vicinity of “Tehuantepec”; this locality does not appear to be the usual collecting site of Tehuantepec, Oaxaca, but rather a community in the municipality of San Andrés Tuxtla called Tehuantepec (from the notes of C.M. Bogert; Pérez-Higareda & Smith 1991). Independently of this, the morphological characteristics of the holotype of bogerti correspond with the morphological characteristics of a population of specimens of Salvadora bairdi collected in the state of Veracruz, near Cordoba. It is worth mentioning that Salvadora bairdi is the species that varies the most in squamation and coloration patterns. After an extensive review of the literature and after many years of collecting (either personal or by other collectors) in the region of Los Tuxtlas, no more specimens have come to light, and it is apparent that the holotype of S. bogerti may have been a mislabeled specimen. Pérez-Higareda et al. (2007) do not list this species as part of the ophidiofauna of the region; therefore, we conclude that the locality of the type of Salvadora bogerti is not from Los Tuxtlas and more likely was collected by Sumichrast near Orizaba; a locality where he lived and made collections. Some individuals in northern Sinaloa, southern Sonora and Chihuahua have a series of patches or transverse patches on the first third of body; however, in spite of being an isolated population, this character is not constant in the specimens we examined.

Salvadora hexalepis (Cope, 1866) Fig. 9 Phimothyra hexalepis Cope, 1866: 304. Subspecies included S. hexalepis hexalepis S. hexalepis virgultea S. hexalepis mojavesis S. hexalepis klauberi Diagnosis Well-developed rostral scale with free lateral edges; normally 9 supralabials, or occasionally 8 or 10; supralabials in contact with the eye, usually only sixth is in contact; 10–11 infralabials; preocular divided; loreal scale normally divided; prenasal scale separated from the second supralabial; a second pair of chinshields separated by two rows of scales; ventral scales 180–213; subcaudal scales vary from 75 to 103; maxillary teeth 10–12 +3. With the exception of S. hexalepis virgultea, the dark dorsolateral lines are normally separated from the lateral lines at the level of the third and fourth rows of dorsal scales; tail length is 20 to 26% of the total body length. Salvadora hexalepis differs at the species level from the other species in the grahamiae group by having a divided preocular, fewer than 113 subcaudals, and a tail length less than 29% of the total length. It differs from S. grahamiae, S. intermedia, S. gymnorhachis and S. bairdi by having 9 to 10 supralabials, the sixth in contact with the eye versus 8 supralabials present in the other species in the grahamiae group. It differs from S. deserticola by having a divided loreal and 75–103 subcaudals. Type material examined UNITED STATES • Arizona, Fort Whipple; USNM 7894. Distribution and conservation This species occurs in the southwestern United States from Nevada, southern Utah, southward through California, Arizona, the Baja California Peninsula and Sonora including Tiburón, San José, and Espíritu Santo Islands in the Gulf of California. (Fig. 9). Within its distribution, S. hexalepis has been recorded in several protected natural areas in Mexico and the United States. Mexico and the United States do not extend legal protection to this species. It is listed by the IUCN as a species of Least Concern., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 99-100, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Cope E. D. 1866. On the Reptilia and Batrachia of the Sonoran Province of the Nearctic region. Proceedings of the Academy of Natural Sciences of Philadelphia 18: 300 - 314."]}

Salvadora lineata Schmidt, 1940 Fig. 8 Salvadora lineata Schmidt, 1940: 143–150. Salvadora grahamiae lineata Hartweg, 1940: 259. Diagnosis Well-developed rostral scale, elongated and with free edges; 8 or 9 supralabials, fourth and fifth in contact with the eye; 9 to 11 infralabials; preocular divided; a single loreal scale; prenasal scale in contact with or separated from the second supralabial; a second pair of chinshields in contact with each other or separated by a row of scales; 179–202 ventral scales; 81–107 subcaudal scales; maxillary teeth normally 10 +3. Color pattern consists of a pale vertebral line reaching the top of the head, three to five rows of scales wide on the first third of the body and only three posteriorly; head bordered by a pair of continuous dorsolateral lines from the loreal region across the body, dark line on the sixth and seventh dorsal scales; lateral line paler than the well-developed dorsolateral line on the third row of dorsal scales. Differs from S. grahamiae by having a pair of well-defined lateral lines on the third row of the dorsal scales, extending from the first third of the body and to the cloaca. Etymology Derived from the Latin ‘ linea ’, meaning ‘line’, in reference to the conspicuous lateral line. Type material examined UNITED STATES • ♀; Texas, Kleberg County, Kingsville; 1938; J.C. Cross; FMNH 28605. Distribution and conservation United States from eastern Texas southward through northwestern Mexico in the states of Chihuahua, Coahuila, Nuevo León, Tamaulipas, Durango, Zacatecas, San Luis Potosí, Guanajuato Querétaro, Michoacán, Hidalgo, Puebla and northern Veracruz (Fig. 8). This species occurs in several protected natural areas in Mexico and the United States. Mexico and the United States have not listed this species in any protection category, but it is on the IUCN red list as a species of Least Concern., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 98-99, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Schmidt K. P. 1940. Notes on Texan snakes of the genus Salvadora. Field Museum of Natural History Zoological Series 24: 143 - 150. https: // doi. org / 10.5962 / bhl. title. 3249","Hartweg N. 1940. Description of Salvadora intermedia, new species, with remarks on the grahamiae group. Copeia 1940 (4): 256 - 259. https: // doi. org / 10.2307 / 1438583"]}

Salvadora mexicana (Duméril, Bibron & Duméril, 1854) Fig. 4 Zamenis mexicanus Duméril, Bibron & Duméril, 1854: 695–696. Salvadora mexicana – Günther 1863: 349. Diagnosis Rostral elongated; 9 supralabials, 3 supralabials in contact with the eye, the fourth, fifth and sixth; 10–11 infralabials; a single preocular; a single loreal scale; prenasal in contact with the second supralabial; a second pair of chinshields in contact with each other or separated by a row of scales; 182–197 ventrals; 125–145 subcaudals; tail length is 31 to 42% of total length; 15 +3 maxillary teeth. Color pattern on the first third of the body consists of mottled transverse patches becoming transverse bars on the neck; the back of the head is dark with pale longitudinal patch in the center; the rest of the body has pale vertebral line flanked by two dark dorsolateral lines; and two dark lateral lines. This species differs from its congeners by having a single preocular scale; tail length is approximately one third of the total length (31–42%); and by having four dark longitudinal lines interrupted on the first third of the body where they are replaced by transverse bands or patches. Etymology Patronym referring to the country of origin. Type material MEXICO • “Cap Corrientes, Mexique” [Jalisco, Cabo Corrientes]; MNHN 3585. Distribution and conservation Endemic to Mexico. Occurs from Nayarit southward on the Pacific coast in the states of Nayarit, Jalisco, Colima, Michoacán, Guerrero, and Oaxaca entering the Balsas River Basin, toward central Guerrero, the states of Estado de México, Morelos and southwestern Puebla (Fig. 4). Within its distribution it is found in at least four protected natural areas. Mexican legislation has listed this species as subject to Special Protection and the IUCN has it listed as a species of Least Concern., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 92-93, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Dumeril A. M., Bibron C. G. & Dumeril A. 1854. Erpetologie generale ou Histoire naturelle complete des Reptiles. Vol. 7 (partie 1). Librairie encyclopedique de Roret. Paris. https: // doi. org / 10.5962 / bhl. title. 118790","Gunther A. 1863. Third account of new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History 12: 348 - 365. https: // doi. org / 10.1080 / 00222936308681536"]}

Salvadora hexalepis mojavensis Bogert, 1935 Fig. 9 Salvadora hexalepis mojavensis Bogert, 1945: 14. Diagnosis Nine supralabials, no supralabial in contact with the eye or only sixth in contact; 10–11 infralabials; from two to three suboculars; 190–207 ventral scales; 82–96 subcaudal scales; tail 20 to 25% of total length; 11+3 maxillary teeth; a pale vertebral line that does not reach the parietal region, bordered by a pair of dorsolateral lines and a pair of somewhat inconspicuous lateral lines and irregular bars along the body that interrupt the dorsolateral and lateral lines. Differs from all conspecific subspecies by having two or three suboculars present. Etymology Toponym alluding to its distribution that is concentrated in the Mojave Desert. Type material examined UNITED STATES • ♂; California, San Bernardino County, Deadman Point, 11.5 miles SE of Victorville, at the south end of the Granite Mountains; 19 Jun. 1935; C.M. Bogert leg; AMNH 63000. Distribution Endemic to the United States. Distributed south of the San Joaquín Valley and the Mojave Desert, east and north of the foothills of the San Gabriel and San Bernardino mountains, to Pyramid Lake in western Nevada, the Virgin River Basin in southern to western Utah, and portions of the Painted Desert in northern Arizona near the Grand Canyon and the canyon of the small Colorado River (Fig. 9)., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on page 102, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Bogert C. M. 1935. Salvadora grahamiae virgultea, a new subspecies of the patch-nosed snake. Bulletin of the Southern California Academy of Sciences 34: 88 - 94.","Bogert C. M. 1945. Two additional races of the patch-nosed snake Salvadora hexalepis. American Museum Novitates 1285: 1 - 14."]}

Salvadora hexalepis virgultea (Bogert, 1935) Fig. 9 Salvadora grahamiae virgultea Bogert, 1935: 89. Diagnosis Nine supralabials, the sixth or the fifth and sixth in contact with the eye; 10–11 infralabials; suboculars absent; 188–200 ventral scales; 84–103 subcaudal scales; tail 21 to 26% of the total length; maxillary teeth 10+3. A pale vertebral line three rows of scales wide is in contact with the parietal region, bordered by a pair of dorsolateral lines brown in color, five rows of scales wide extending from the temporal region to the tip of the tail. At about midbody the lateral line gives rise to a thinner lateral line that is one row of scales wide. Salvadora h. klauberi differs in having 12 + 3 maxillary teeth; in S. h. mojavensis suboculars separate the supralabials from the lower edge of the eye; S. h. hexalepis sometimes also lacks suboculars and has a single supralabial in contact with the eye similar to S. h. virgultea, but it differs in its color pattern (in S. h. virgultea the vertebral line involves three scale rows; the head is brown, and a lateral line is not distinguishable). In S. h. hexalepis the vertebral line is up to five scale rows wide; the head is gray; and the lateral line at midbody is completely separated from the dorsolateral line. Etymology From the Latin ‘ virgultus ’, meaning ‘covered with scrub’, for the habitat in which the taxon occurs. Type material examined UNITED STATES • ♂; California, San Diego County; 32.67778° N, 116.72709° W; 29 Jun. 1929; Deerhorn Flat leg.; SDSNH 12025. Distribution Regions adjacent to the Pacific coast and adjacent foothills in southwestern California, southward to the southern portion of Baja California, where is it parapatric with S. h. kaluberi (Fig. 9)., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 101-102, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Bogert C. M. 1935. Salvadora grahamiae virgultea, a new subspecies of the patch-nosed snake. Bulletin of the Southern California Academy of Sciences 34: 88 - 94.","Cope E. D. 1866. On the Reptilia and Batrachia of the Sonoran Province of the Nearctic region. Proceedings of the Academy of Natural Sciences of Philadelphia 18: 300 - 314.","Bogert C. M. 1945. Two additional races of the patch-nosed snake Salvadora hexalepis. American Museum Novitates 1285: 1 - 14."]}

Salvadora hexalepis klauberi Bogert, 1945 Fig. 9 Salvadora hexalepis klauberi Bogert, 1945: 14. Diagnosis Nine supralabials, generally fifth and sixth in contact with the eye; 10–11 infralabials; suboculars normally absent; 180–203 ventral scales; 78–99 subcaudal scales; tail 21 to 26% of total length; 12 +3 maxillary teeth; a pale vertebral line three to five rows wide contacting the parietal region, bordered by a pair of dark dorsolateral lines up to four rows wide extending from the temporal region to the margins between the fifth and fourth rows of scales, lateral lines one or two scales wide fused with the dorsolateral lines on first third of the body. Differs from S. h. mojavensis by lacking suboculars, differs from S. h. hexalepis by having fifth and sixth supralabials bordering the eye and differs from S. h. virgultea by having 12 +3 maxillary teeth. Etymology Dedicated to Laurence Monroe Klauber. Type material MEXICO • ♂; Baja California Sur, Cape San Lucas; 6 Aug. 1933; Fred Lewis leg.; Collection of L.M. Klauber No. 20912. Distribution Endemic to Mexico, present from northern Baja California southward through Baja California Sur, including the San José and Espíritu Santo Islands (Fig. 9)., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 102-103, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Bogert C. M. 1945. Two additional races of the patch-nosed snake Salvadora hexalepis. American Museum Novitates 1285: 1 - 14."]}

Salvadora gymnorhachis Hernández, Flores & Campbell, 2019 Fig. 11 Diagnosis Eight–9 supralabials, two supralabials, fourth and fifth in contact with the eye; 8–10 infralabials; preocular scale divided; a single loreal scale; prenasal in contact or separated from the second supralabial; a second pair of chinshields in contact or separated by a row of scales; 176–186 ventral scales; 92–103 subcaudal scales; tail is 18 to 21% of the total length; 11 +3 maxillary teeth. Color pattern consists of a pair of dark dorsolateral lines that begin behind the eyes and fade before reaching the middle of the body, without a well-defined vertebral line and lacking lateral lines. This species differs clearly from the other species by lacking dorsolateral lines on about the posterior two-thirds of the body and by lacking vertebral and lateral lines across the body. Etymology The name is derived from the Greek prefixes ‘ gymnós ’, which means ‘naked’ and ‘ ráchḭ ’ which means ‘back’. Type material examined MEXICO • ♀; Oaxaca, Distrito Mixe, San Pedro and San Pedro Ayutla; 17.00159° N, 96.08443° W; 24 Aug. 2014; MZFC 28775. Distribution and conservation Mountains of central Oaxaca, Mexico (Fig.11). This is a recently described taxon and one of limited distribution that is not known to inhabit any protected natural areas. This species is not included in Mexican legislation and is not in the IUCN’s red list. Dichotomous key for the species and subspecies of the genus Salvadora 1. A single preocular (Fig. 12A); more than 123 subcaudals; three supralabials in contact with the eye; maxillary teeth 14 + 3 to 15 +3; tail length more than 29% of total body length (mexicana group). 2 – Preocular generally divided (Fig. 12B); fewer than 116 subcaudals; normally fewer than three supralabials in contact with the eye; maxillary teeth from 9+3 to 12 +3; tail length generally less than 29% of total body length (grahamiae group)............................................................................ 3 2. Four longitudinal lines on the posterior part of the body that are interrupted in the first third of the body by transverse patches or crossbars (Fig. 13A); 182–197 ventral scales; tail length 31% to 42% of total length; maxillary teeth 15 + 3.................... S. mexicana (Duméril, Bibron & Duméril, 1854) – Four well-defined longitudinal lines extending along the body without transverse patches or crossbars on the first third of body (Fig. 13B); 194 to 205 ventral scales; tail length between 29% and 33% of total length; maxillary teeth 14 +3........................................................... S. lemniscate (Cope, 1895) 3. Supralabials normally eight (rarely 7 or 9), fourth and fifth in contact with the eye (Fig. 14A); a second pair of chinshields in contact with each other or separated by one row of scales (Fig. 15A........................................................................................................................................................... 4 – Nine or ten supralabials (rarely 11 or 12), generally the sixth is in contact with the eye (Fig. 14B); posterior pair of chinshields sometimes separated by two rows of scales (Fig.15B)........................ 8 4. Maxillary teeth 11 +3; dorsolateral lines normally present along the body or incomplete present only in the first third of body, lateral line normally present on third or fourth row of dorsal scales, or absent when the dorsolateral line is incomplete (populations from Guerrero, Oaxaca or south of Puebla)............................................................................................................................................... 5 – Maxillary teeth 9 +3 or 10 +3; dorsolateral line always present; lateral line well defined on the third or fourth row of dorsal scales, or vestigial or absent......................................................................... 6 5. Dorsolateral lines present along the entire body; a light, yellowish vertebral line; lateral lines always present; tail length 23 to 30 % of total body length.............................. S. intermedia Hartweg, 1940 – Dorsolateral lines absent on the second and last third of the body; no lateral lines on body; tail length is 19 to 21 % of total length.......................................................................................................................................................... S. gymnorhachis Hernández-Jiménez, Flores-Villela & Campbell, 2019 6. Maxillary teeth 9+ 3; vertebral line normally separated from the parietal region by a patch formed by the fusion of the dorsolateral lines (Fig. 16A); dorsolateral lines do not reach the temporal region; vertebral line three scale rows wide on the first and second third of the body (sometimes interrupted by dark blotches), and one row wide on the last third; lateral line well defined and separated from the dorsolateral line the entire length of the body............................... S. bairdi Jan & Sordelli, 1860 – Maxillary teeth 10 + 3; vertebral line reaching the parietal region (Fig. 16B); dorsolateral lines pass through the temporal region to the eye; vertebral line from three to five rows wide on the first third of the body, and three rows wide on the second and last third of the body; lateral line present, vestigial or barely discernible, or may be absent in some specimens............................................................. 7 7. Lateral line absent, vestigial, or scarcely discernible (Fig. 17A)............................................................................................................................................................ S. grahamiae Baird & Girard, 1853 – Lateral line always present and well defined (Fig. 17B).............................. S. lineata Schmidt, 1940 8. Loreal scale normally single (Fig. 18A); vertebral line five rows wide on the first third of body; lateral line well defined, largely restricted to adjoining margins of scales in third and fourth rows, clearly separated from the dorsolateral line on the second and last thirds of body; 68 to 86 subcaudals.............................................................................................. S. deserticola Schmidt, 1940 – Loreal normally divided (Fig. 18B); vertebral line generally 3 rows wide, sometimes more; lateral line ill-defined or well-defined but fused or feebly separated from the dorsolateral lines; 75 to 103 subcaudals.......................................................................................................................................... 9 9. Vertebral line yellowish in color, no more than three scale rows in width along the entire body; dorsolateral lines brown fused with the lateral lines in the anterior part of body but feebly separated posteriorly, generally without subocular scales, overall appearance brown......................................................................................................................................................... S. h. virgultea Bogert,1935 – Vertebral line light grey in color, more than three rows wide (3 to 7) on the first third of the body; dorsolateral lines brown or grey in color, and four rows wide on the first third of the body, three rows wide in the second and one row wide on the last third of the body, and regularly separated or slightly separated from the lateral lines, with or without subocular scales.................................................. 10 10. Normally two supralabials reaching the eye (Fig. 19A); 12 +3 maxillary teeth.................................................................................................................................................. S. h. klauberi Bogert,1945 – Fewer than two supralabial reaching the eye (Fig. 19B); 10 or 11 +3 maxillary teeth....................11 11. Supralabials normally separated from the eye by the presence of two or more subocular scales (Fig. 20A);10–11 infralabial scales; 11 +3 maxillary teeth; sometimes with an irregular pattern of bars running the length of the body that interrupt the dorsolateral and lateral lines......................................................................................................................................... S. h. mojavensis Bogert,1945 – One supralabial in contact with the eye (Fig. 20B); normally without suboculars or when present there is only one; normally 10 infralabials; 10+3 maxillary teeth; vertebral line three scales in width; dorsolateral line nearly gray, often irregularly pigmented, and obscurely separated from lateral line anteriorly, over-all appearance gray....................................................... S. h. hexalepis (Cope, 1866), Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 105-111, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Hernandez-Jimenez C. A., Flores-Villela O. & Campbell J. A. 2019. A new species of patch-nosed snake (Colubridae: Salvadora Baird and Girard, 1853) from Oaxaca, Mexico. Zootaxa 4564 (2): 588 - 600. https: // doi. org / 10.11646 / zootaxa. 4564.2.14","Dumeril A. M., Bibron C. G. & Dumeril A. 1854. Erpetologie generale ou Histoire naturelle complete des Reptiles. Vol. 7 (partie 1). Librairie encyclopedique de Roret. Paris. https: // doi. org / 10.5962 / bhl. title. 118790","Cope E. D. 1895. The classification of the Ophidia. Transactions of the American Philosophical Society 18: 186 - 219. https: // doi. org / 10.2307 / 1005387","Hartweg N. 1940. Description of Salvadora intermedia, new species, with remarks on the grahamiae group. Copeia 1940 (4): 256 - 259. https: // doi. org / 10.2307 / 1438583","Jan G. & Sordelli F. 1860. Iconographie generale des Ophidiens. 1. Livraison. J. B. Bailiere et Fils, Paris. https: // doi. org / 10.5962 / bhl. title. 45246","Baird S. F. & Girard C. 1853. Catalogue of North American Reptiles in the Museum of the Smithsonian Institution. Part 1 Serpents. Smithsonian Institution, Washington D. C. https: // doi. org / 10.5962 / bhl. title. 5513","Schmidt K. P. 1940. Notes on Texan snakes of the genus Salvadora. Field Museum of Natural History Zoological Series 24: 143 - 150. https: // doi. org / 10.5962 / bhl. title. 3249","Bogert C. M. 1935. Salvadora grahamiae virgultea, a new subspecies of the patch-nosed snake. Bulletin of the Southern California Academy of Sciences 34: 88 - 94.","Bogert C. M. 1945. Two additional races of the patch-nosed snake Salvadora hexalepis. American Museum Novitates 1285: 1 - 14.","Cope E. D. 1866. On the Reptilia and Batrachia of the Sonoran Province of the Nearctic region. Proceedings of the Academy of Natural Sciences of Philadelphia 18: 300 - 314."]}

The grahamiae group The taxa in this group are defined by several synapomorphies, including preocular generally divided; fewer than 116 subcaudals, normally fewer than three supralabials in contact with the eye, maxillary teeth from 9 to 12 +3; tail length less than 29% of total length., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on page 94, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543

Salvadora hexalepis hexalepis (Cope, 1866) Fig. 9 Diagnosis Nine supralabials; sixth supralabial in contact with the eye; 10 infralabials; a single subocular sometimes present; 186–213 ventrals; 75–98 subcaudals; tail 20 to 25 % of total length; 10 +3 maxillary teeth; a pale vertebral line contacting the parietal region, three to five scale rows wide on the first third of the body, three rows wide posteriorly, bordered by a pair of dorsolateral lines up to four scale rows wide, extending from the temporal region to the tip of the tail, halfway along the body the dorsolateral lines are divided giving rise to another lateral line distinguishable to the tip of tail. This taxon differs from S. h. klauberi by having a single supralabial in contact with the eye, whereas S. h. klauberi has two supralabials (the fifth and sixth) in contact; in S. h. mojavensis the supralabials and the lower edge of the eyes are separated by two or more suboculars; S. h. hexalepis is similar to S. h. virgultea in that many individuals have a single supralabial in contact with the eye; however, these subspecies differ in coloration pattern, with the head of S. h. virgultea being brown and the lateral line becoming faded and indistinguishable at about the middle of the body (in S. h. hexalepis the head is gray, the vertebral line is up to five rows of scales wide, and in the middle of the body the lateral line is completely separated from the dorsolateral line). Etymology From the Latin ‘ hexa ’ for ‘six’ and ‘ lepis ’ for ‘scale’, which can be interpreted as six-sided scales. Distribution It occurs from the southeast of California, United States, southward to northeastern Baja California, central Arizona southward through the northeast portion of the Sonoran Desert, including Tiburón Island (Fig. 9)., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 100-101, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Cope E. D. 1866. On the Reptilia and Batrachia of the Sonoran Province of the Nearctic region. Proceedings of the Academy of Natural Sciences of Philadelphia 18: 300 - 314."]}

Salvadora grahamiae Baird & Girard, 1853 Fig. 7 Salvadora grahamiae Baird & Girard, 1853: 104–105. Phimothyra grahamiae – Cope 1861: 566. Diagnosis Well-developed rostral scale, elongated and with free edges; 8 or 9 supralabials, fourth and fifth or fifth and sixth supralabial in contact with the eye; normally 10 infralabials; a single loreal scale; preocular divided; prenasal scale in contact with the second supralabial; a second pair of chinshields separated by a row of scales; 178–200 ventral scales and 89–106 subcaudals; maxillary teeth normally 10 +3. Color pattern consists of a pale vertebral line extending to the top of the head, three to five rows of scales wide on the first third of the body reduced to only three on posteriorly; head bordered by a pair of continuous dorsolateral lines from the loreal region throughout the body, with dark line on the sixth and seventh lines of dorsal scales; Sometimes with an inconspicuous lateral line at third row of dorsal scales. Salvadora grahamiae differs from S. lineata by having inconspicuous or absent lateral lines, and differs from other congeners by having the tail approximately one quarter of the total length (22–29%); loreal scale not divided, posterior chinshields separated by a two row of scales; and the vertebral line up to five scales wide on anterior of body. Etymology Dedicated to the collector of the taxon, J.D. Graham. Type material examined UNITED STATES • ♂; Arizona, Cochise County, Huachuca Mountains, limited to Sonora (Bogert 1939a); J.D. Graham leg.; USNM 2081. Distribution and conservation From western Texas, northern Coahuila, Chihuahua, and Sonora to northern Arizona and New Mexico (Fig. 7). It occurs in several protected natural areas in Mexico and the United States. Mexico and the United States have not listed this species in any protection category, and it is on the IUCN red list as a species of Least Concern., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on page 97, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Baird S. F. & Girard C. 1853. Catalogue of North American Reptiles in the Museum of the Smithsonian Institution. Part 1 Serpents. Smithsonian Institution, Washington D. C. https: // doi. org / 10.5962 / bhl. title. 5513","Cope E. D. 1861. Catalogue of the Colubridae in the Museum of the Academy of Natural Sciences of Philadelphia. Part 3. Proceedings of the Academy of Natural Sciences of Philadelphia 12: 553 - 566.","Bogert C. M. 1939 a. A study of the genus Salvadora. The patch-nosed snakes. Publications of the University of California at Los Angeles, Biological Sciences 1: 177 - 236."]}

Salvadora lemniscata (Cope, 1895) Fig. 3 Drymobius lemniscatus Cope, 1895: 203. Drymobius (Eudryas) pulcherrimus Bocourt 1890: 725–726. Salvadora lemniscata – Bogert 1939b: 142. Diagnosis Rostral slightly elongated and projecting toward the back; 9 supralabials, 3 supralabials in contact with the eye, the fourth, fifth and sixth; generally 12 infralabials, sometimes 11 or 13; single preocular; single loreal; prenasal separated from the second supralabial; a second pair of chinshields larger than the first one in contact with each other or separated by a row of scales; 194–206 ventrals; 128–141 subcaudals; tail is 29 to 34% of total length; 14–15+3 maxillary teeth. Color pattern consists of a pale vertebral line extending from the parietal region, and is three rows wide on the first third of the body and one row wide on the rest of the body, flanked by a pair of dark dorsolateral lines two to four scales wide, interrupted at the neck without touching the head; two dark lateral lines, sometimes slightly interrupted in the neck region or continuing toward the sides of the head, passing by the eyes to the nasal scales. Differs from the other species in the genus by having more maxillary teeth (14–15 +3); a single preocular scale; tail length 29 to 34% of total length; 4 longitudinal lines along the body, dorsolaterals interrupted on the neck region; 194 to 206 ventral scales. Etymology The species name comes from the Latin ‘ lemniscatus ’, which means ‘adorned with bands’, doubtless in reference to the stripes along its body. Type material MEXICO • “Mexique occidental” [Western Mexico], locality uncertain, probably from Tehuantepec, Oaxaca; F. Sumichrast leg.; MNHN 1891.0260. Distribution and conservation From Acapulco in the state of Guerrero southward the Pacific coast, across the Isthmus of Tehuantepec to the coast of Chiapas, the Central Depression of the Grijalva River, and Huehuetenango, Guatemala (Fig. 3). Within its distribution it occurs in at least two protected natural areas. Mexican legislation has listed this species as subject to Special Protection and the IUCN has it listed as a species of Least Concern. Remarks Three additional specimens examined (UTA 45840 Guatemala: Huehuetenango, 2.4 km from Nenton on Road to la Trinidad; UTA 45842 Guatemala: Huehuetenango, 0.3 km S Cuatro caminos, La Democracia- Nenton Road; UTA 45843 Guatemala Nenton, Hacienda Miramar, La Fortuna near Pozo) represent the first records of Salvadora lemniscata for Guatemala., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on page 91, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Cope E. D. 1895. The classification of the Ophidia. Transactions of the American Philosophical Society 18: 186 - 219. https: // doi. org / 10.2307 / 1005387","Bocourt M. F. 1890. Mission scientifique au Mexique dans l'Amerique centrale. Reptiles. 12: 697 - 732.","Bogert C. M. 1939 b. Notes on snakes of the genus Salvadora with a redescription of a neglected Mexican species. Copeia 1939 (3): 140 - 147. https: // doi. org / 10.2307 / 1436808"]}

Salvadora deserticola Schmidt 1940 Fig. 6 Salvadora hexalepis deserticola Schmidt, 1940: 146, fig. 14. Salvadora hexalepis celeris Smith, 1941: 9, fig. 6. Diagnosis Rostral scale well developed, elongated and with free edges; 9 supralabials, two or rarely three in contact with the eye (generally the fifth and sixth in contact); 11 infralabials; preocular scale divided; a single loreal; prenasal in contact with the second supralabial; a second pair of chinshields separated by two rows of scales; 180–205 ventrals; 66–87 subcaudals; tail is 19 to 23.5% of the total length; maxillary teeth normally 11+ 3. Color pattern consists of a pale vertebral stripe, five scales wide on the anterior third of the body and three scales wide on the rest of the body, flanked by a pair of dark dorsolateral lines located on the sixth and seventh row and separated from the lateral lines by at least one row of scales; lateral lines begin on the fourth row or on the margins of the third and fourth rows of dorsal scales. Lateral and dorsolateral lines fused on the neck. Differs from the other species in the genus by having a single loreal, usually not divided; tail shorter (less than 24% of the total length) and fewer subcaudal scales. Etymology The species name is derived from the Latin words ‘ desertum ’, which means ‘desert’, and ‘ icola ’, which means ‘inhabitant’, in reference to the habitat at the type locality. Type material examined UNITED STATES • ♂, holotype of Salvadora hexalepis deserticola; Texas, Brewster County, Government Spring, close to the Chisos Mountains; 6 Oct. 1935; Tom Carney leg.; FMNH 26615. MEXICO • ♀, holotype of Salvadora hexalepis celeris; Sinaloa, San Blas; 28 Mar. 1910; J.N. Rose leg.; USNM 40043. Distribution and conservation Southwestern Texas, southwestern New Mexico, and southeastern Arizona, southward on each flank of the Sierra Madre Occidental in Mexico in the states of Sonora, Chihuahua, Coahuila and Sinaloa (Fig. 6). It is found in at least two protected natural areas in Sonora. Mexican legislation has not listed this species and the IUCN has it listed as a species of Least Concern., Published as part of Hernández-Jiménez, Carlos A., Flores-Villela, Oscar, Aguilar-Bremauntz, Aranzazú & Campbell, Jonathan A., 2021, Phylogenetic relationships based on morphological data and taxonomy of the genus Salvadora Baird & Girard, 1853 (Reptilia, Colubridae), pp. 85-118 in European Journal of Taxonomy 764 (1) on pages 95-97, DOI: 10.5852/ejt.2021.764.1473, http://zenodo.org/record/5514543, {"references":["Schmidt K. P. 1940. Notes on Texan snakes of the genus Salvadora. Field Museum of Natural History Zoological Series 24: 143 - 150. https: // doi. org / 10.5962 / bhl. title. 3249","Smith H. M. 1941. Further notes on Mexican snakes of the genus Salvadora. Smithsonian Miscellaneous Collections 99: 1 - 12. Available from https: // repository. si. edu / handle / 10088 / 23709 [accessed 23 Jul. 2021]."]}

Salvadora gymnorhachis, sp. nov. Figs. 1, 2 urn:lsid:zoobank.org:act: FA3E5069-814D-4890-BC45-489F9AF4133B Paratypes. Five, all from the state of Oaxaca, Mexico. A juvenile female (AMNH 84992) from San Pablo Mitla, 1900 m; an adult male (AMNH 91090) from 8.3 km north of Mitla, 2100 m; and an adult female (AMNH 100910) from 2 km west of San Lorenzo Albarradas, 2000 m; all three from Distrito de Tlacolula. An adult female (AMNH 100909) from 12 km (via road) southwest of San Pedro y San Pablo Ayutla, 1760 m, Distrito de Mixe; and an adult female (UIMNH 56849), near Santa Catalina Quier��, 2364 m, Distrito de Yautepec. Diagnosis. Salvadora gymnorhachis differs from all its congeners by lacking a pale vertebral line, and dorsolateral stripes extending only to about the middle of the body (vs other species in which these stripes extend by throughout whole length of the body) (Table 1); Salvadora gymnorhachis also differs from S. grahamiae, S. bairdi, S. mexicana, and S. lemniscata by having 11 maxillary teeth followed by a diastema and three enlarged teeth (9���10+ 3 in S. grahamiae and S. bairdi; 14���15+ 3 in S. mexicana and S. lemniscata); it further differs from S. mexicana and S. lemniscata by having fewer than 105 subcaudals (vs more than 124), a divided preocular (vs single in S. mexicana and S. lemniscata); from S. deserticola and S. hexalepis by having second and third supralabials in contact with loreal (vs third, fourth, and sometimes fifth contacting loreal); and from S. intermedia by having six and five scale rows separating dorsolateral stripes on anterior and middle part of body, respectively (vs 3 and 1-2 scale rows separating stripes on anterior and middle part of body, respectively). Description of holotype. Rostral with free lateral edges, extending slightly between internasal suture; plates behind rostral on top of head including prefrontals, supraoculars, frontal and paired parietals; on side of head prenasal separated from second supralabial; loreal situated above second and third supralabials; two preoculars, upper larger, contacting prefrontal and supraocular; lower preocular small, situated above third and fourth supralabials; two postoculars at left side, lower contacting fifth and sixth supralabials; one postocular at right side touching the large fifth supralabial; temporals 2 + 2, lower contacting fifth, sixth, and seventh supralabials (Fig. 2A); 8/8 supralabials; fourth and fifth touching eye; 8/8 infralabials, first pair meeting at midventral line behind small mental, first four pairs reaching anterior genials; fourth infralabial enlarged more than length of anterior genials; posterior genials smaller than anterior genials, separated by single row of two median scales; dorsal scales in 19 rows immediately behind head at level of ventral 3, 17 rows at level of ventral scale 13; 16 rows at midbody at level of ventral scale 90, and 13 scale rows anterior to vent at level of ventral scale 170; 182 ventrals; cloacal scute divided, bordered laterally by small scales; tail incomplete, 80+ subcaudals; 11 maxillary teeth, increasing in size posteriorly, followed by diastema and three enlarged teeth (11+3). Measurements. Total length 612 mm; SVL 471 mm; TaL 141 mm (tip missing), comprising about 23 % of total length. Coloration. Pattern consisting of a pair of dark dorsolateral stripes (89 Jet black; Smithe, 1975) beginning at level of postoculars, extending to middle of body where stripes grade into ground color; dorsolateral stripes occupying upper half of second dorsal scale row and covering four scale rows on anterior part of body, reduced to three scales wide just before becoming inconspicuous at about mid-body; field between dorsolateral stripes brown (223B Verona brown), six rows wide at anterior part of body, five at middle of body where stripes begin to fade; posterior part of body gray (45 Smoke gray); ventral surface of body whitish (92 Pale horn) except for salmon color (6 Salmon) on lateral portion of ventrals of posterior part of body and venter of tail; lateral edges of ventral scales black. Variation of Paratypes. Except for minor differences in scutelation, paratypes closely resemble holotype (Table 2); ventrals in females 176���184, single male with 186 ventrals; subcaudals 92���103 in females, tail incomplete in single male; supralabials 8/ 8 in most specimens, except in female from near Municipality of San Lorenzo Albarradas, Oaxaca with 9 on one side; third and fourth supralabial reaching second (lower) postocular on individuals from District of Tlacolula, Oaxaca, whereas female from Municipality of Santa Catalina Quieri, Oaxaca (UIMNH 56849) and female from Municipality of San Pedro y San Pablo Ayutla, Oaxaca (AMNH R��� 100909) with only fifth supralabial touching postocular; anterior nasal in contact with second supralabial in all specimens, except the one from Municipality of Mitla (AMNH 91090), Oaxaca, in which anterior nasal slightly separated from second supralabials; two supralabials, reaching loreal in all specimens except for one female (AMNH 100910) with three on left side; body length 852 to 975 mm, male being largest in series and female holotype smallest (Table 2). Male closely resembling females, including holotype, in lacking vertebral stripe; male paler, however, with little more than vestiges of dorsolateral stripes; prior to preservation lips and throat white, in contrast to yellow venter of anterior portion of trunk and two lowermost rows of scales (Bogert, unpublished notes); posteriorly yellow color of the venter grading into salmon color extending to underside of tail; basal portions of lower dorsal scales on the posterior of trunk appreciably darker than margins; individual having brownish body flanked with yellow anteriorly gradually becoming gray speckled with black posteriorly; dorsolateral stripes obsolescent in preserved specimen, more conspicuous in live snake. Etymology. The specific epithet comes from the Greek ��� gymn��s ���, meaning ���naked���, and ��� r��ch��� ��� dorsum, in reference that this is the only species of Salvadora that does not have dorsal vertebral stripes and has incomplete dorsolateral ones. Habitat and Conservation. The holotype was found beneath a rock in a transitional area between a crop field and pine-oak forest (Fig. 3A). The distribution of this species undoubtedly has been greatly modified by anthropogenic activities, which has increased the rate of deforestation at the type locality (Fig. 3B). None of the collection sites are located in protected areas. Despite intensive fieldwork in Oaxaca, only six specimens of this species have been collected, all between 1960 and 2014. Distribution. Most individuals have been collected in central Oaxaca along the unnumbered State secondary road between Mitla and Ayutla in the Distritos de Tlacolula and Mixe at elevations of 1760 to 2364 m. The species is also known in the south near Santa Catalina Quier�� in Distrito de Yautepec at elevations exceeding 1600 m. The entire range of this species is drained by the upper tributaries of the R��o Tehuantepec (Fig. 4). Most of the river valleys of the region contain fingers of upper tropical deciduous forest extending from lower coastal regions and the higher slopes and ridges are covered with pine-oak forest. The phylogenetic analysis performed gave one phylogenetic tree (Fig. 5) with maximun likelihood value of - 2709.59. The genus Salvadora is retrieved as monophyletic, with sampled species forming two strongly supported clades (mexicana group and grahamiae group). Salvadora gymnorachis is nested within the grahamiae group as the sister taxon of Salvadora intermedia., Published as part of Hern��ndez-Jim��nez, Carlos Alberto, Flores-Villela, Oscar & Campbell, Jonathan Atwood, 2019, of patch-nosed snake (Colubridae: Salvadora Baird and Girard, 1853) from Oaxaca, Mexico, pp. 588-600 in Zootaxa 4564 (2) on pages 589-596, DOI: 10.11646/zootaxa.4564.2.14, http://zenodo.org/record/2589083, {"references":["Smithe, F. B. (1975) Naturalist Color Guide. American Museum of Natural History, New York, 229 pp."]}

Salvadora gymnorhachis, sp. nov. Figs. 1, 2 urn:lsid:zoobank.org:act: FA3E5069-814D-4890-BC45-489F9AF4133B Paratypes. Five, all from the state of Oaxaca, Mexico. A juvenile female (AMNH 84992) from San Pablo Mitla, 1900 m; an adult male (AMNH 91090) from 8.3 km north of Mitla, 2100 m; and an adult female (AMNH 100910) from 2 km west of San Lorenzo Albarradas, 2000 m; all three from Distrito de Tlacolula. An adult female (AMNH 100909) from 12 km (via road) southwest of San Pedro y San Pablo Ayutla, 1760 m, Distrito de Mixe; and an adult female (UIMNH 56849), near Santa Catalina Quierí, 2364 m, Distrito de Yautepec. Diagnosis. Salvadora gymnorhachis differs from all its congeners by lacking a pale vertebral line, and dorsolateral stripes extending only to about the middle of the body (vs other species in which these stripes extend by throughout whole length of the body) (Table 1); Salvadora gymnorhachis also differs from S. grahamiae, S. bairdi, S. mexicana, and S. lemniscata by having 11 maxillary teeth followed by a diastema and three enlarged teeth (9–10+ 3 in S. grahamiae and S. bairdi; 14–15+ 3 in S. mexicana and S. lemniscata); it further differs from S. mexicana and S. lemniscata by having fewer than 105 subcaudals (vs more than 124), a divided preocular (vs single in S. mexicana and S. lemniscata); from S. deserticola and S. hexalepis by having second and third supralabials in contact with loreal (vs third, fourth, and sometimes fifth contacting loreal); and from S. intermedia by having six and five scale rows separating dorsolateral stripes on anterior and middle part of body, respectively (vs 3 and 1-2 scale rows separating stripes on anterior and middle part of body, respectively). Description of holotype. Rostral with free lateral edges, extending slightly between internasal suture; plates behind rostral on top of head including prefrontals, supraoculars, frontal and paired parietals; on side of head prenasal separated from second supralabial; loreal situated above second and third supralabials; two preoculars, upper larger, contacting prefrontal and supraocular; lower preocular small, situated above third and fourth supralabials; two postoculars at left side, lower contacting fifth and sixth supralabials; one postocular at right side touching the large fifth supralabial; temporals 2 + 2, lower contacting fifth, sixth, and seventh supralabials (Fig. 2A); 8/8 supralabials; fourth and fifth touching eye; 8/8 infralabials, first pair meeting at midventral line behind small mental, first four pairs reaching anterior genials; fourth infralabial enlarged more than length of anterior genials; posterior genials smaller than anterior genials, separated by single row of two median scales; dorsal scales in 19 rows immediately behind head at level of ventral 3, 17 rows at level of ventral scale 13; 16 rows at midbody at level of ventral scale 90, and 13 scale rows anterior to vent at level of ventral scale 170; 182 ventrals; cloacal scute divided, bordered laterally by small scales; tail incomplete, 80+ subcaudals; 11 maxillary teeth, increasing in size posteriorly, followed by diastema and three enlarged teeth (11+3). Measurements. Total length 612 mm; SVL 471 mm; TaL 141 mm (tip missing), comprising about 23 % of total length. Coloration. Pattern consisting of a pair of dark dorsolateral stripes (89 Jet black; Smithe, 1975) beginning at level of postoculars, extending to middle of body where stripes grade into ground color; dorsolateral stripes occupying upper half of second dorsal scale row and covering four scale rows on anterior part of body, reduced to three scales wide just before becoming inconspicuous at about mid-body; field between dorsolateral stripes brown (223B Verona brown), six rows wide at anterior part of body, five at middle of body where stripes begin to fade; posterior part of body gray (45 Smoke gray); ventral surface of body whitish (92 Pale horn) except for salmon color (6 Salmon) on lateral portion of ventrals of posterior part of body and venter of tail; lateral edges of ventral scales black. Variation of Paratypes. Except for minor differences in scutelation, paratypes closely resemble holotype (Table 2); ventrals in females 176–184, single male with 186 ventrals; subcaudals 92–103 in females, tail incomplete in single male; supralabials 8/ 8 in most specimens, except in female from near Municipality of San Lorenzo Albarradas, Oaxaca with 9 on one side; third and fourth supralabial reaching second (lower) postocular on individuals from District of Tlacolula, Oaxaca, whereas female from Municipality of Santa Catalina Quieri, Oaxaca (UIMNH 56849) and female from Municipality of San Pedro y San Pablo Ayutla, Oaxaca (AMNH R– 100909) with only fifth supralabial touching postocular; anterior nasal in contact with second supralabial in all specimens, except the one from Municipality of Mitla (AMNH 91090), Oaxaca, in which anterior nasal slightly separated from second supralabials; two supralabials, reaching loreal in all specimens except for one female (AMNH 100910) with three on left side; body length 852 to 975 mm, male being largest in series and female holotype smallest (Table 2). Male closely resembling females, including holotype, in lacking vertebral stripe; male paler, however, with little more than vestiges of dorsolateral stripes; prior to preservation lips and throat white, in contrast to yellow venter of anterior portion of trunk and two lowermost rows of scales (Bogert, unpublished notes); posteriorly yellow color of the venter grading into salmon color extending to underside of tail; basal portions of lower dorsal scales on the posterior of trunk appreciably darker than margins; individual having brownish body flanked with yellow anteriorly gradually becoming gray speckled with black posteriorly; dorsolateral stripes obsolescent in preserved specimen, more conspicuous in live snake. Etymology. The specific epithet comes from the Greek “ gymnós ”, meaning “naked”, and “ ráchḭ ” dorsum, in reference that this is the only species of Salvadora that does not have dorsal vertebral stripes and has incomplete dorsolateral ones. Habitat and Conservation. The holotype was found beneath a rock in a transitional area between a crop field and pine-oak forest (Fig. 3A). The distribution of this species undoubtedly has been greatly modified by anthropogenic activities, which has increased the rate of deforestation at the type locality (Fig. 3B). None of the collection sites are located in protected areas. Despite intensive fieldwork in Oaxaca, only six specimens of this species have been collected, all between 1960 and 2014. Distribution. Most individuals have been collected in central Oaxaca along the unnumbered State secondary road between Mitla and Ayutla in the Distritos de Tlacolula and Mixe at elevations of 1760 to 2364 m. The species is also known in the south near Santa Catalina Quierí in Distrito de Yautepec at elevations exceeding 1600 m. The entire range of this species is drained by the upper tributaries of the Río Tehuantepec (Fig. 4). Most of the river valleys of the region contain fingers of upper tropical deciduous forest extending from lower coastal regions and the higher slopes and ridges are covered with pine-oak forest. The phylogenetic analysis performed gave one phylogenetic tree (Fig. 5) with maximun likelihood value of - 2709.59. The genus Salvadora is retrieved as monophyletic, with sampled species forming two strongly supported clades (mexicana group and grahamiae group). Salvadora gymnorachis is nested within the grahamiae group as the sister taxon of Salvadora intermedia.

Phyllodactylus ruPinus Dixon, 1964 Holotype. Adult male. KU 67501, collected by James R. Dixon. Mike Sabath, and Richard Worthington. from 7 miles south Lombardia, on the bank of the Rio Marquez. Michoacan. Elevation 461 meters, August 25, 1960. Diagnosis. A medium-sized gecko although on average larger than P. kropotkini; it presents 28 tubercles from head to tail, similar to P. lupitae (28.8) and P. kropotkini (28.4). It differs from P. isabelae (32.2), P. lanei (33.2), P. benedettii (27.4), P. t. magnus (37), P. muralis (33.6) and P. tuberculosus (36.4) in tubercular counts. Phyllodactylus rupinus presents 14 rows of tubercles across its dorsum similar to P. t. magnus (14.3) and P. tuberculosus (14), and differs from P. isabelae (16.7), P. lupitae (14.8), P. lanei (15.6), P. benedettii (13.8), P. kropotkini (13.4) and P. muralis (12.2). Phyllodactylus rupinus is clearly differentiated from other gecko species because it presents 26.5 scales across venter, while the others present: P. isabelae (27.8), P. lupitae (24.6), P. lanei (30), P. benedettii (29.1), P. kropotkini (30.6), P. t. magnus (27.33), P. muralis (33) and P. tuberculosus (30.2). Distribution. This species is restricted to the locality of Lombardia, Michoacan., Published as part of Ram��rez-Reyes, Tonatiuh & Flores-Villela, Oscar, 2018, Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico, pp. 151-190 in Zootaxa 4407 (2) on page 175, DOI: 10.11646/zootaxa.4407.2.1, http://zenodo.org/record/1216693, {"references":["Dixon, J. R. (1964) The systematics and distribution of lizards of the genus Phyllodactylus in North and Central America. New Mexico State University Research Center, Scientific Bulletin, 1964, 1 - 139."]}

Phyllodactylus benedettii sp. nov. Holotype. Adult male (MZFC 28774) collected at Chamela Biological Station, La Huerta, state of Jalisco (19.50 N, - 105.44W WGS84) on September 15, 2014 by Tonatiuh Ram��rez Reyes and Alfredo Villarruel. Paratypes (14). An adult female (MZFC 28773) collected at Chamela Biology Station, La Huerta municipality (19.50 N, - 105.44W) on September 15, 2014 by Tonatiuh Ram��rez Reyes and Alfredo Villarruel; (MZFC 21817- male, MZFC 21818-male, MZFC 21877, MZFC 21878-male, MZFC 21879, MZFC 21880-female, MZFC 21885- male) collected at the Chamela Biology Station, La Huerta municipality (19.50 N, - 105.44 W) by Oscar Flores Villela in 2007; an adult female (IBH 2133-2) and five adult males (IBH 2133-3, 2133-4, 2133-5, 2133-7, 2133-9) collected at 2 km. SE, of the Station of Biology, Chamela (19.51 N, - 104.94 W) on October 27, 1971 by Cornelio S��nchez; an adult female (IBH 2138-4) collected at 5 km. S., de Chamela, Biology Station, UNAM, municipality La Huerta (19.49 N, - 104.95 W) on May 25, 1974 by Gustavo Casas Andreu. Etymology. The species is dedicated to the memory of the great Uruguayan writer Mario Benedetti in recognition of his prolific literary production and critical thinking of great importance in the political and social life of Latin America. Diagnosis. Phyllodactylus benedettii sp. nov. is a species with the largest average size within the P. lanei complex, with a maximum SVL of 75.2 mm. Dixon (1964) established three fundamental characteristics to include species and subspecies within the P. lanei complex and differentiate them from P. tuberculosus: species of the P. lanei complex have a low number of interorbital scales (12���19), white venter and low number of scales across the snout between the third labials (does not specify range of values). Phyllodactylus benedettii shows white-yellow venter, 14.4 interorbital scales on average, and 22.3 scales across the snout between the third labials. Within the P. lanei complex (including insular and continental species), the only species that exceed 70 mm of SVL are P. lupitae and P. lanei, two species that are very isolated geographically. It is clearly different from the nearest species P. rupinus, which has a maximum SVL of 69.4 mm, considered as a medium size species. According to the statistical analyzes previously carried out, the combination of diagnostic morphological characters are the following (in addition to the SVL): It differs clearly from all species of Phyllodactylus as it presents 62.6 (60���69) longitudinal scales, 27 (24���30) tubercles from head to tail, 14 (11���13) interorbital scales, 29 (25���32) scales across venter, 22 (22���23) third labial���snout scales, 13 (13���14) rows of tubercles across dorsum. Although P. benedettii is very similar to P. lupitae in some meristic characters, they present differences in the morphometric space (Fig. 7). Some measures allow us to differentiate these species are the head-length and axillagroin length: P. benedettii (17.9 mm and 28.4 mm) and P. lupitae (16.86 mm and 27.93 mm). According to Castiglia et al. (2009; 2010), the karyotypes of P. l. lanei (Tierra Colorada, Guerrero) and P. l. rupinus from Chamela, Jalisco (here named P. benedetti) are different. According to their studies P. lanei has 2n = 33���34 and FN = 40���41, while P. benedettii has 2n = 38 and FN = 38. Phyllodactylus benedettii has fewer than 30 tubercles from head to tail (24���30, 27.4 avg.) similar to P. lupitae (27���31, 28.8 avg.), P. paucituberculatus (28.7), P. kropotkini (25���31, 28.4 avg.) and P. rupinus (26���30, 28 avg.); other species have more than 30 tubercles from head to tail on average: P. lanei (32���34, 33.2 avg.), P. isabelae (30��� 35, 32.2 avg.), P. t. magnus (35���40, 37 avg.), P. muralis (30���40, 33 avg.) and P. tuberculosus (33���41, 36.4 avg.). Phyllodactylus benedettii showed 62 longitudinal ventral scales (LVS) on average (60���69), other values were P. isabelae (52���58, 56.2 avg.), P. lupitae (60���64, 61.8 avg.), P. lanei (60���74, 66 avg.), P. rupinus (56���67, 63 avg.), P. kropotkini (63���73, 67 avg.), P. t. magnus (52���58, 54 avg.), P. muralis (57���61, 59 avg.) and P. tuberculosus (51���64, 57 avg.). Phyllodactylus benedettii presents 14.4 interorbital scales similar to P. kropotkini (14���16, 14.6 avg.) and differs from P. isabelae (14���20, 15.1 avg.), P. lupitae (14���20, 16.8 avg.), P. lanei (15���17, 15.5 avg.), P. rupinus (14���17, 15.5 avg.), P. t. magnus (21���23, 23.3 avg.), P. muralis (21���27, 23.2 avg.), and P. tuberculosus (16���22, 19.2 avg.). Phyllodactylus benedettii presents 29 scales across venter and differs from P. isabelae (26���29, 27.8 avg.), P. lupitae (26���29, 24.6 avg.), P. lanei (29���32, 30 avg.), P. rupinus (24���28, 26.5 avg.), P. kropotkini (29���33, 30.6 avg.), P. t. magnus (26���29, 27.3 avg.), P. muralis (33) and P. tuberculosus (27���33, 30.2 avg.). Phyllodactylus benedettii presents 22 third labial���snout scales on average (22���23) and differs from P. isabelae (19���24, 21.4 avg.), P. lupitae (23���28, 25.5 avg.), P. lanei (20���23, 21.2 avg.), P. rupinus (18���22, 20.6 avg.), P. kropotkini (19���21, 20.2 avg.), P. t. magnus (24���26, 24.6 avg.), P. muralis (24���28, 25.2 avg.) and P. tuberculosus (21���26, 24.2 avg.). Phyllodactylus benedettii presents 13.8 rows of tubercles across dorsum on average (13���14) and differs from P. isabelae (15���18, 16.7 avg.), P. lupitae (14���15, 14.8 avg.), P. lanei (14���16, 15.6 avg.), P. rupinus (13���15, 14 avg.), P. kropotkini (12��� 14, 13.4 avg.), P. t. magnus (13���15, 14.3 avg.), P. muralis (12���13, 12.2 avg.) and P. tuberculosus (12���17, 14 avg.). Finally, P. benedettii presents the largest values in the following measures: 5.50 mm in length of the 4th toepad and 35.8 mm in axilla-groin length (LAG); compared to P. isabelae (4.1 mm, 22.2 mm), P. lupitae (4.9 mm, 31 mm), P. lanei (4.6 mm, 21.1 mm), P. rupinus (5 mm, 25.6 mm) and P. kropotkini (5 mm, 23 mm). Description of holotype (Fig. 9). (All bilateral counts are given as right/left). Adult male, robust body, head not flattened, neck slightly differentiated from the head. The head scales are granular, most are slightly globose, and are interspersed with circular-oblique flat scales that are mainly located between the eyes towards the tip of the snout. The rostral scale presents an intermediate striation near to middle of the scale (a groove that extends longitudinally to the middle part of the rostral scale); nostril bordered by a simple supranasal (one supranasal scale only), first labial, rostral and two postnasals; two supranasal scales contact each other, bordered by 7 postnasals; 22 interorbital scales beginning at the anterior ocular border; 15 scales across the snout from the second right labial, 24 scales across the snout at the level of the third labial scale on the right side; 8/9 loreal scales from the first ocular scale to the nostril; 11/10 supralabial scales; posterior border of the first supralabial in contact with the nostril (right and left); 8/8 infralabial scales; elongated auricular opening smaller than ocular diameter; occipital scales different in size and shape to the interorbital scales, while the interorbital scales are more or less uniform in size and shape (oval shape), they are smaller than the occipital ones. The occipital scales are larger than the interorbital scales, they have irregular shapes and presents some circular tubercles intermixed; mental scale is equally wide as long (3.6 mm), forms an irregular "V" towards the posterior section, two postmental scales closely in contact each other and with the first and second infralabials; postmental scales in contact with a row of 4 scales, the 2 intermediates larger than the extremes, 10 scales of uniform size border the row of scales. Body with granular scales; 13 rows of strongly keeled dorsal tubercles with variable size; 25 tubercles in a middle dorsal row from base of the head to base of the tail; 21/15 tubercles from axilla to groin; 34 rows of cycloid scales across the venter; 87 longitudinal ventral scales from the first differentiated scale on the gular region to the cloacal opening; ventral scales larger compared to the lateral scales of the body and the gular region; scales slightly imbricate on the anterior and posterior extremities, scales dorsally granular intercalated with strongly keeled tubercles of variable size. The tail of the specimen is original (not regenerated) and measures 55.5 mm with the tip broken. Dorsal scales of the tail are imbricate with intercalated tubercles and juxtaposed ventral scales, keeled tubercles extend to more than half of the tail, as they extend to the tip of the tail the tubercles gradually flatten until they form keeled scales. Does not present femoral or precloacal pores. Digital lamella formulae: right posterior (7-9-12-13-10), left posterior (6-8-11-13-9); right anterior (6-9-12-11- 11), left anterior (6-10-13-11-12), fourth finger of the extremities longer than others; digital toepads longer than wide on all fingers. Measurements in mm: snout-vent length 70.9; axilla-groin 35.8; head length 16.5; head width 13.2; snout length (to eye) 10.05; eye diameter 4.1; auricular opening (maximum) 1.6; length of fourth finger 5.5; length of fourth toe 5.7; width between eye supercilliaries 9; internaral 2.1. Coloration of holotype in ethanol (Fig. 8). The holotype exhibits relatively little coloration, the dorsal background color varies from brown to creamy yellow, with dark brown faint irregular spots (less evident compared to live coloration), the ventral surface varies yellowish and cream; half of the ventral color on the tail to the tip is cream. Pattern and color in life (Fig. 10). Dorsally presents a clear background, a mixture of cream-pink, on this background, patterns of irregular patches dark brown color and some completely black spots are present, some of these spots are slightly surrounded by yellowish scales; on the middle dorsal part a longitudinal line of the background color runs from head to tail; scales around the eyes and some supralabials are slightly colored light yellow; ventrally presents a white-cream coloration in combination with light yellow, more evident towards the ventral region; the pupils are black on a bronze iris. Dorsal coloration of the tail is white-cream with brown stripes and some black points. Variation. All meristic and morphometric characters are presented with mean values, standard deviation and range of meristic (Table 2) and morphometric variables (Table 7). Distribution and habitat. Phyllodactylus benedettii sp. nov. is an endemic species to the state of Jalisco, restricted to the Chamela-Cuixmala Biosphere Reserve. According to the INEGI digital map of elevation (resolution of 15 m), the type locality is located at 100 masl, located in the biogeographic province of the Western Pacific surrounded by deciduous tropical forests (Fig. 11). The microhabitats reported for this species are diverse: soil, trees, shrubs and in habitats modified by man as houses or bridges or some other constructions (meteorological stations). They are particularly abundant in rocks and crevices and in people's houses or buildings where it feeds on insects and other invertebrates (Garc��a & Ceballos 1994; Ponce-Campos & Garc��a 2007; Vitt & Caldwell 2014; pers. obs.). Males of P. benedettii have an extended period of reproduction (August���March) with two peaks, one in rainy season (August���October) and another one in dry season (November���March). Reproductive behavior (courtship and mating) begins in August which coincides with the beginning of rainy season. The females produce clutches of two eggs, with up to three clutches during the breeding season, with a peak of egg production from December to March coinciding with the dry season (Ram��rez-Sandoval et al. 2006). This species presents a distinct karyotype from P. lanei presumably due to Robertsonian fusions/fissions with 19 pairs of telocentric chromosomes (2n = 38, FN = 38) (Castiglia et al. 2009). They are lizards of nocturnal habits and feed on insects, although to date there have been no studies on the composition of their diet (Garc��a & Ceballos 1994; Vitt & Caldwell 2014). Representative species of tropical dry forest of Chamela-Cuixmala Biosphere Reserve are: Sciadodendrom excelsum, Brosimum alicastrum, Orbignya cohune and Tabebuia donell-smithi (Ceballos et al. 1999). The deciduous tropical forests is near the mouth of the Cuixmala River and is characterized by the abundance of water in the rainy season, the climate is tropical (warm-humid), has an annual average temperature of 25 ��C and the annual rainfall varies from 748���1000 mm (Garc��a & Ceballos 1994; Ceballos et al. 1999). Other species of reptiles with nocturnal activity that inhabit the deciduous tropical forests are: Hemidactylus frenatus, Boa constrictor, Lampropeltis triangulum, Leptodeira maculata, Pseudoficimia frontalis, Senticolis triaspis, Tropidodipsas philippii (Sibon philippi), Tantilla bocourti, Thamnophis valida, Trimorphodon biscutatus, Micrurus distans, Agkistrodon bilineatus, and Crotalus basiliscus (Garc��a & Ceballos 1994). Comments on conservation and threats. This species inhabits the Chamela-Cuixmala Biosphere Reserve decreed in 1993. There is a minor threat posed by the localities bordering the reserve, and according to the observations made by the first author, apparently there is a large population of P. benedettii geckos inside Chamela- Cuixmala Biology Station (UNAM). There are two main threats: the false beliefs of some people who consider this species as poisonous (local inhabitants), and the possible negative effects due to the presence of invasive gecko Hemidactylus frenatus, a species with a very large population that may exploit the same ecological niche. To date there is no specific study on niche overlap between H. frenatus and Phyllodactylus species in Mexico, however, its potential danger to local biodiversity has been documented, including interspecific negative interactions with native geckos from different areas of the world; one example of this is the displacement and reduction of six gecko species of the genus Nactus on the Mascarene Islands and Lepidodactylus lugubris throughout Pacific islands. In these cases H. frenatus has been observed stalking, lunging towards and biting at other geckos (Global Invasive Species Database 2015). Due to the above, and as a potential risk to local biodiversity in Mexico, it is classified as a high risk invasive species (CONABIO 2016). Phyllodactylus kroPotkini sp. nov. Holotype. Adult male (MZFC 28736) collected in Nueva Filadelfia (Huerta Vieja) in the municipality of Tlapehuala, Guerrero (18.29 N, - 100.49 W WGS84), collected on March 15, 2014 by Tonatiuh Ram��rez Reyes and Joel Rosas Avila. Paratypes (4). Four male adults (MZFC 28735, MZFC 28737, MZFC 28738, MZFC 28739) collected at the type locality, Nueva Filadelfia (Huerta Vieja), Tlapehuala, Guerrero (18.29 N, - 100.49 W WGS84) collected on March 16, 2014 by Tonatiuh Ram��rez Reyes and Joel Rosas ��vila. Etymology. The species is dedicated to the memory of the great Russian philosopher, geographer and naturalist Piotr Kropotkin, who made great scientific and theoretical contributions about mutual support and altruism in some animal populations (including human society). Diagnosis. Phyllodactylus kropotkini sp. nov. is a medium-sized gecko within the P. lanei complex. Dixon (1964) established three fundamental characteristics to include species and subspecies within the P. lanei complex and differentiate them from P. tuberculosus, species of the P. lanei complex have a low number of interorbital scales (12���19), white venter and low number of scales across the snout between the third labials (does not specify range of values). Phyllodactylus kropotkini has 14.6 interorbital scales on average (14���15), white venter and 20.2 third labial���snout scales on average (19���21). Phyllodactylus kropotkini presents a maximum recorded SVL of 62.3 mm that clearly differs from the nearest species P. lanei (max. 71 mm) and it differs from other species: P. isabelae (max. 57.5), P. lupitae (max. 73.3 mm), P. rupinus (max. 69.4), P. benedettii (max. 74.2) and P. bordai (max. 58.9 mm). Phyllodactylus kropotkini sp. nov. shows the following combination of characters: 28 (25���31) tubercles from head to tail, 67 (63���73) longitudinal scales, 13 (12���14) rows of tubercles across dorsum, 30 (29���33) scales across venter, 14 (14���15) interorbital scales and 20 (19���21) third labial���snout scales. Phyllodactylus kropotkini has a partial overlap with P. lanei and P. isabelae on morphometric space (Fig. 7), however P. kropotkini is clearly differentiated from P. lanei by maximum length (SVL) (Fig. 6); it also presents the follow measures: axilla-groin length (22.54 mm), snout length (9.34 mm), auricular opening (1.78 mm), while P. lanei has 24.7 mm axilla-groin length, 10. 61 mm snout length and auricular opening (1.95 mm) and P. isabelae has 18.7 mm axilla-groin length, 8.18 mm snout length and auricular opening 1.51 mm. P. kropotkini has a low number of interorbital scales (14���15, 14.6 avg.) similar to P. benedettii (13���16, 14.4 avg.), all other species of Phyllodactylus exceed 15 interorbital scales on average, P. isabelae (14���16, 15.1 avg.), P. lupitae (14���20, 16.83 avg.), P. lanei (15���17, 15.5 avg.), P. rupinus (14���17, 15.5 avg.), P. t. magnus (21���23, 22.3 avg.), P. muralis (21���27, 23.2 avg.), P. tuberculosus (16���22, 19.2 avg.). There are an average of 30.6 scales across the venter (29���33), unlike P. isabelae (26���29, 27.8 avg.), P. lupitae (23���26, 24.6 avg.), P. lanei (29���32, 30 avg.), P. rupinus (24���28, 26.5 avg.) and P. benedettii (25���32, 29.1 avg.). Phyllodactylus kropotkini presents 67.6 longitudinal scales from the gular region to the anus on average (63���73), compared to P. rupinus (63���73, 63.6 avg.), P. lanei (60���74, 66 avg.), P. benedettii (60���69, 62.6 avg.), P. t. magnus (52���58, 54.6 avg.), P. mura, Published as part of Ram��rez-Reyes, Tonatiuh & Flores-Villela, Oscar, 2018, Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico, pp. 151-190 in Zootaxa 4407 (2) on pages 168-174, DOI: 10.11646/zootaxa.4407.2.1, http://zenodo.org/record/1216693, {"references":["Dixon, J. R. (1964) The systematics and distribution of lizards of the genus Phyllodactylus in North and Central America. New Mexico State University Research Center, Scientific Bulletin, 1964, 1 - 139.","Castiglia, R, Aguayo, A. G., Bezerra, A. M. R., Flores-Villela, O. & Gournung, E. (2009) Karyotypic diversification due to Robertsonian rearrangements in Phyllodactylus lanei Smith, 1935 (Squamata, Gekkonidae) from Mexico. Atti della Accademia Nazionale dei Lincei, Rendiconti Lincei, Scienze Fisiche e Naturali, 20, 77 - 82. https: // doi. org / 10.1007 / s 12210 - 009 - 0005 - 4","Castiglia, R., Annesi, F., Bezerra, A. M. R., Garcia, A. & Flores-Villela, O. (2010) Cytotaxonomy and DNA taxonomy of lizards (Squamata, Sauria) from a tropical dry forest in the Chamela-Cuixmala Biosphere Reserve on the coast of Jalisco, Mexico. Zootaxa, 2508, 1 - 29.","Garcia, A. & Ceballos, G. (1994) Guia de campo de los reptiles y anfibios de la costa de Jalisco, Mexico. Fundacion Ecologica de Cuixmala, A. C., Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico, D. F., 176 pp.","Ponce-Campos, P. & Garcia-Aguayo, A. (2007) Phyllodactylus lanei. The IUCN Red List of Threatened Species. Version 2014.2. Available from: http: // www. iucnredlist. org (accessed 27 October 2014)","Vitt, L. J. & Caldwell, J. P. (2014) Herpetology: An Introductory Biology of Amphibians and Reptiles. 4 th Edition. Academic Press, San Diego, 757 pp. https: // doi. org / 10.1016 / B 978 - 0 - 12 - 386919 - 7.00002 - 2","Ramirez-Sandoval, E., Ramirez-Bautista, A. & Vitt, J. L. (2006) Reproduction in the lizard Phyllodactylus lanei (Squamata: Gekkonidae) from the Pacific Coast of Mexico. Copeia, 1, 1 - 9. https: // doi. org / 10.1643 / 0045 - 8511 (2006) 006 [0001: RITLPL] 2.0. CO; 2","Ceballos, G., Szekely, A., Garcia, A., Rodriguez, P. & Noguera, F. (1999) Programa de Manejo de la Reserva de la Biosfera Chamela-Cuixmala. Instituto Nacional de Ecologia, SEMARNAP, Mexico, D. F., 133 pp.","CONABIO (2016) Sistema de informacion sobre especies invasoras en Mexico. Comision Nacional para el Conocimiento y Uso de la Biodiversidad. Available from: http: // www. biodiversidad. gob. mx / invasoras (accessed 13 March 2018)","Toledo, A. (2003) Rios, costas, mares. Hacia un analisis integrado de las regiones hidrologicas de Mexico. INE-Semarnat, Mexico, 117 pp.","Pineda-Garcia, F., Arredondo-Amezcua, L. & Ibarra-Manriquez, G. (2007) Riqueza y diversidad de especies lenosas del bosque tropical caducifolio El Tarimo, Cuenca del Balsas, Guerrero. Revista Mexicana de Biodiversidad, 78, 129 - 139.","Reyna-Alvarez, J., Suazo-Ortuno, I. & Alvarado-Diaz, J. (2010) Herpetofauna del Municipio de Huetamo, Michoacan, Mexico. Biologicas, 12 (1), 40 ¯ 45.","Smith, H. M. (1935) Miscellaneous notes on Mexican lizards. The University of Kansas Science Bulletin, 36, 119 - 155.","Castro-Franco, R. & Uribe-Pena, Z. (1992) Dos subespecies nuevas de Phyllodactylus lanei (Sauria: Gekkonidae). Anales del Instituto de Biologia. Universidad Nacional Autonoma de Mexico, Serie Zoologia, 63, 113 - 123."]}

Phyllodactylus lanei Smith, 1935 Holotype. Adult male, CNHM 100067, collected by Edward H. Taylor and Hobart M. Smith near Tierra Colorada Guerrero, June 30, 1932. Diagnosis. A large gecko according to Dixon (1964) attaining a snout-vent length of 78 mm. This observation agrees with ours, as the largest species in the P. lanei complex are P. lanei, P. benedettii and P. lupitae. Has more than 30 tubercles from head to tail (33.2), similar to P. isabelae (32.2) and P. muralis (33.6). The other species of the P. lanei complex have fewer than 30, P. lupitae (28.8), P. rupinus (28), P. benedettii (27.4), P. kropotkini (28.4). Other species of geckos with more tubercles are P. t. magnus (37) and P. tuberculosus (36.4). Phyllodactylus lanei presents 30 scales across the venter similar to P. kropotkini (30.6) and P. tuberculosus (30.2), but differing from P. isabelae (27.8), P. lupitae (24.6), P. rupinus (26.5), P. benedettii (29.1) and P. t. magnus (27.3). Distribution. This species is restricted to the localities of Tierra Colorada and Chilpancingo, Guerrero and adjacent locations of medium-low altitude localities., Published as part of Ram��rez-Reyes, Tonatiuh & Flores-Villela, Oscar, 2018, Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico, pp. 151-190 in Zootaxa 4407 (2) on page 175, DOI: 10.11646/zootaxa.4407.2.1, http://zenodo.org/record/1216693, {"references":["Smith, H. M. (1935) Miscellaneous notes on Mexican lizards. The University of Kansas Science Bulletin, 36, 119 - 155.","Dixon, J. R. (1964) The systematics and distribution of lizards of the genus Phyllodactylus in North and Central America. New Mexico State University Research Center, Scientific Bulletin, 1964, 1 - 139."]}

Phyllodactylus luPitae Castro-Franco & Uribe-Pe��a, 1992 Holotype. Adult female, EBUM 1242 collected by R. Gonzalo Gavi��o de la Torre on La Pe��a Island (or Coral island), Nayarit, Mexico, November 21, 1986. Diagnosis. This is a larger gecko along with P. lanei and P. benedettii of the P. lanei complex. It presents 28.8 tubercles from head to tail, similar to P. rupinus (28) and P. kropotkini (28.4). These counts differ from the other gecko species: P. isabelae (32.2), P. lanei (33.2), P. benedettii (27.4), P. t. magnus (37), P. muralis (33.6) and P. tuberculosus (36.4). Phyllodactylus lupitae differs clearly from the other species as it shows 25.5 third labial���snout scales, the others: P. isabelae (21.4), P. lanei (21.2), P. rupinus (20.6), P. benedettii (22.3), P. kropotkini (20.2), P. t. magnus (24.6), P. muralis (25.2) and P. tuberculosus (24.2). It is a unique species because it shows 24.6 scales across venter, the others: P. isabelae (27.8), P. lanei (30), P. rupinus (26.5), P. benedettii (29.1), P. kropotkini (30.6), P. t. magnus (27.3), P. muralis (33) and P. tuberculosus (30.2). Also presents 61.8 longitudinal scales, the others: P. isabelae (56.2), P. lanei (66), P. rupinus (63.6), P. benedettii (62.6), P. kropotkini (67.6), P. t. magnus (54.6), P. muralis (59) and P. tuberculosus (57.7). Distribution. This species is endemic to Coral Island (or La Pe��a island), Nayarit, Mexico., Published as part of Ram��rez-Reyes, Tonatiuh & Flores-Villela, Oscar, 2018, Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico, pp. 151-190 in Zootaxa 4407 (2) on pages 175-176, DOI: 10.11646/zootaxa.4407.2.1, http://zenodo.org/record/1216693, {"references":["Castro-Franco, R. & Uribe-Pena, Z. (1992) Dos subespecies nuevas de Phyllodactylus lanei (Sauria: Gekkonidae). Anales del Instituto de Biologia. Universidad Nacional Autonoma de Mexico, Serie Zoologia, 63, 113 - 123."]}

Phyllodactylus isabelae Castro-Franco & Uribe-Peña, 1992 Holotype. An adult female, ZUP 1152 collected by Zeferino Uribe Peña on Marietas Islands, Nayarit, Mexico, February 23, 1978. Diagnosis. This species clearly differs from the others because it is the smallest of the analyzed in the present study (Fig. 6). It presents 56.2 longitudinal scales on average, it differs clearly from the others: P. lupitae (61.8), P. lanei (66), P. rupinus (63.6), P. benedettii (62.6), P. kropotkini (67.6), P. t. magnus (54.6), P. muralis (59) and P. tuberculosus (57.7). Also P. isabelae presents 27.8 scales across the venter similar to P. t. magnus (27.3) and differs from the others: P. lupitae (24.6), P. lanei (30), P. rupinus (26.5), P. benedettii (29.1), P. kropotkini (30.6), P. muralis (33) and P. tuberculosus (30.2). Phyllodactylus isabelae differs clearly from the others analyzed since it has the highest number of rows of tubercles across dorsum (16.7 on average) compared to the others: P. lupitae (14.8), P. lanei (15.6), P. rupinus (14), P. benedettii (13.8), P. kropotkini (13.4), P. t. magnus (14.3), P. muralis (12.2) and P. tuberculosus (14). Distribution. Phyllodactylus isabelae is endemic of the Marietas islands, Nayarit, Mexico.

Phyllodactylus isabelae Castro-Franco & Uribe-Pe��a, 1992 Holotype. An adult female, ZUP 1152 collected by Zeferino Uribe Pe��a on Marietas Islands, Nayarit, Mexico, February 23, 1978. Diagnosis. This species clearly differs from the others because it is the smallest of the analyzed in the present study (Fig. 6). It presents 56.2 longitudinal scales on average, it differs clearly from the others: P. lupitae (61.8), P. lanei (66), P. rupinus (63.6), P. benedettii (62.6), P. kropotkini (67.6), P. t. magnus (54.6), P. muralis (59) and P. tuberculosus (57.7). Also P. isabelae presents 27.8 scales across the venter similar to P. t. magnus (27.3) and differs from the others: P. lupitae (24.6), P. lanei (30), P. rupinus (26.5), P. benedettii (29.1), P. kropotkini (30.6), P. muralis (33) and P. tuberculosus (30.2). Phyllodactylus isabelae differs clearly from the others analyzed since it has the highest number of rows of tubercles across dorsum (16.7 on average) compared to the others: P. lupitae (14.8), P. lanei (15.6), P. rupinus (14), P. benedettii (13.8), P. kropotkini (13.4), P. t. magnus (14.3), P. muralis (12.2) and P. tuberculosus (14). Distribution. Phyllodactylus isabelae is endemic of the Marietas islands, Nayarit, Mexico., Published as part of Ram��rez-Reyes, Tonatiuh & Flores-Villela, Oscar, 2018, Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico, pp. 151-190 in Zootaxa 4407 (2) on page 175, DOI: 10.11646/zootaxa.4407.2.1, http://zenodo.org/record/1216693, {"references":["Castro-Franco, R. & Uribe-Pena, Z. (1992) Dos subespecies nuevas de Phyllodactylus lanei (Sauria: Gekkonidae). Anales del Instituto de Biologia. Universidad Nacional Autonoma de Mexico, Serie Zoologia, 63, 113 - 123."]}

Phyllodactylus benedettii sp. nov. Holotype. Adult male (MZFC 28774) collected at Chamela Biological Station, La Huerta, state of Jalisco (19.50 N, - 105.44W WGS84) on September 15, 2014 by Tonatiuh Ramírez Reyes and Alfredo Villarruel. Paratypes (14). An adult female (MZFC 28773) collected at Chamela Biology Station, La Huerta municipality (19.50 N, - 105.44W) on September 15, 2014 by Tonatiuh Ramírez Reyes and Alfredo Villarruel; (MZFC 21817- male, MZFC 21818-male, MZFC 21877, MZFC 21878-male, MZFC 21879, MZFC 21880-female, MZFC 21885- male) collected at the Chamela Biology Station, La Huerta municipality (19.50 N, - 105.44 W) by Oscar Flores Villela in 2007; an adult female (IBH 2133-2) and five adult males (IBH 2133-3, 2133-4, 2133-5, 2133-7, 2133-9) collected at 2 km. SE, of the Station of Biology, Chamela (19.51 N, - 104.94 W) on October 27, 1971 by Cornelio Sánchez; an adult female (IBH 2138-4) collected at 5 km. S., de Chamela, Biology Station, UNAM, municipality La Huerta (19.49 N, - 104.95 W) on May 25, 1974 by Gustavo Casas Andreu. Etymology. The species is dedicated to the memory of the great Uruguayan writer Mario Benedetti in recognition of his prolific literary production and critical thinking of great importance in the political and social life of Latin America. Diagnosis. Phyllodactylus benedettii sp. nov. is a species with the largest average size within the P. lanei complex, with a maximum SVL of 75.2 mm. Dixon (1964) established three fundamental characteristics to include species and subspecies within the P. lanei complex and differentiate them from P. tuberculosus: species of the P. lanei complex have a low number of interorbital scales (12–19), white venter and low number of scales across the snout between the third labials (does not specify range of values). Phyllodactylus benedettii shows white-yellow venter, 14.4 interorbital scales on average, and 22.3 scales across the snout between the third labials. Within the P. lanei complex (including insular and continental species), the only species that exceed 70 mm of SVL are P. lupitae and P. lanei, two species that are very isolated geographically. It is clearly different from the nearest species P. rupinus, which has a maximum SVL of 69.4 mm, considered as a medium size species. According to the statistical analyzes previously carried out, the combination of diagnostic morphological characters are the following (in addition to the SVL): It differs clearly from all species of Phyllodactylus as it presents 62.6 (60–69) longitudinal scales, 27 (24–30) tubercles from head to tail, 14 (11–13) interorbital scales, 29 (25–32) scales across venter, 22 (22–23) third labial–snout scales, 13 (13–14) rows of tubercles across dorsum. Although P. benedettii is very similar to P. lupitae in some meristic characters, they present differences in the morphometric space (Fig. 7). Some measures allow us to differentiate these species are the head-length and axillagroin length: P. benedettii (17.9 mm and 28.4 mm) and P. lupitae (16.86 mm and 27.93 mm). According to Castiglia et al. (2009; 2010), the karyotypes of P. l. lanei (Tierra Colorada, Guerrero) and P. l. rupinus from Chamela, Jalisco (here named P. benedetti) are different. According to their studies P. lanei has 2n = 33–34 and FN = 40–41, while P. benedettii has 2n = 38 and FN = 38. Phyllodactylus benedettii has fewer than 30 tubercles from head to tail (24–30, 27.4 avg.) similar to P. lupitae (27–31, 28.8 avg.), P. paucituberculatus (28.7), P. kropotkini (25–31, 28.4 avg.) and P. rupinus (26–30, 28 avg.); other species have more than 30 tubercles from head to tail on average: P. lanei (32–34, 33.2 avg.), P. isabelae (30– 35, 32.2 avg.), P. t. magnus (35–40, 37 avg.), P. muralis (30–40, 33 avg.) and P. tuberculosus (33–41, 36.4 avg.). Phyllodactylus benedettii showed 62 longitudinal ventral scales (LVS) on average (60–69), other values were P. isabelae (52–58, 56.2 avg.), P. lupitae (60–64, 61.8 avg.), P. lanei (60–74, 66 avg.), P. rupinus (56–67, 63 avg.), P. kropotkini (63–73, 67 avg.), P. t. magnus (52–58, 54 avg.), P. muralis (57–61, 59 avg.) and P. tuberculosus (51–64, 57 avg.). Phyllodactylus benedettii presents 14.4 interorbital scales similar to P. kropotkini (14–16, 14.6 avg.) and differs from P. isabelae (14–20, 15.1 avg.), P. lupitae (14–20, 16.8 avg.), P. lanei (15–17, 15.5 avg.), P. rupinus (14–17, 15.5 avg.), P. t. magnus (21–23, 23.3 avg.), P. muralis (21–27, 23.2 avg.), and P. tuberculosus (16–22, 19.2 avg.). Phyllodactylus benedettii presents 29 scales across venter and differs from P. isabelae (26–29, 27.8 avg.), P. lupitae (26–29, 24.6 avg.), P. lanei (29–32, 30 avg.), P. rupinus (24–28, 26.5 avg.), P. kropotkini (29–33, 30.6 avg.), P. t. magnus (26–29, 27.3 avg.), P. muralis (33) and P. tuberculosus (27–33, 30.2 avg.). Phyllodactylus benedettii presents 22 third labial–snout scales on average (22–23) and differs from P. isabelae (19–24, 21.4 avg.), P. lupitae (23–28, 25.5 avg.), P. lanei (20–23, 21.2 avg.), P. rupinus (18–22, 20.6 avg.), P. kropotkini (19–21, 20.2 avg.), P. t. magnus (24–26, 24.6 avg.), P. muralis (24–28, 25.2 avg.) and P. tuberculosus (21–26, 24.2 avg.). Phyllodactylus benedettii presents 13.8 rows of tubercles across dorsum on average (13–14) and differs from P. isabelae (15–18, 16.7 avg.), P. lupitae (14–15, 14.8 avg.), P. lanei (14–16, 15.6 avg.), P. rupinus (13–15, 14 avg.), P. kropotkini (12– 14, 13.4 avg.), P. t. magnus (13–15, 14.3 avg.), P. muralis (12–13, 12.2 avg.) and P. tuberculosus (12–17, 14 avg.). Finally, P. benedettii presents the largest values in the following measures: 5.50 mm in length of the 4th toepad and 35.8 mm in axilla-groin length (LAG); compared to P. isabelae (4.1 mm, 22.2 mm), P. lupitae (4.9 mm, 31 mm), P. lanei (4.6 mm, 21.1 mm), P. rupinus (5 mm, 25.6 mm) and P. kropotkini (5 mm, 23 mm). Description of holotype (Fig. 9). (All bilateral counts are given as right/left). Adult male, robust body, head not flattened, neck slightly differentiated from the head. The head scales are granular, most are slightly globose, and are interspersed with circular-oblique flat scales that are mainly located between the eyes towards the tip of the snout. The rostral scale presents an intermediate striation near to middle of the scale (a groove that extends longitudinally to the middle part of the rostral scale); nostril bordered by a simple supranasal (one supranasal scale only), first labial, rostral and two postnasals; two supranasal scales contact each other, bordered by 7 postnasals; 22 interorbital scales beginning at the anterior ocular border; 15 scales across the snout from the second right labial, 24 scales across the snout at the level of the third labial scale on the right side; 8/9 loreal scales from the first ocular scale to the nostril; 11/10 supralabial scales; posterior border of the first supralabial in contact with the nostril (right and left); 8/8 infralabial scales; elongated auricular opening smaller than ocular diameter; occipital scales different in size and shape to the interorbital scales, while the interorbital scales are more or less uniform in size and shape (oval shape), they are smaller than the occipital ones. The occipital scales are larger than the interorbital scales, they have irregular shapes and presents some circular tubercles intermixed; mental scale is equally wide as long (3.6 mm), forms an irregular "V" towards the posterior section, two postmental scales closely in contact each other and with the first and second infralabials; postmental scales in contact with a row of 4 scales, the 2 intermediates larger than the extremes, 10 scales of uniform size border the row of scales. Body with granular scales; 13 rows of strongly keeled dorsal tubercles with variable size; 25 tubercles in a middle dorsal row from base of the head to base of the tail; 21/15 tubercles from axilla to groin; 34 rows of cycloid scales across the venter; 87 longitudinal ventral scales from the first differentiated scale on the gular region to the cloacal opening; ventral scales larger compared to the lateral scales of the body and the gular region; scales slightly imbricate on the anterior and posterior extremities, scales dorsally granular intercalated with strongly keeled tubercles of variable size. The tail of the specimen is original (not regenerated) and measures 55.5 mm with the tip broken. Dorsal scales of the tail are imbricate with intercalated tubercles and juxtaposed ventral scales, keeled tubercles extend to more than half of the tail, as they extend to the tip of the tail the tubercles gradually flatten until they form keeled scales. Does not present femoral or precloacal pores. Digital lamella formulae: right posterior (7-9-12-13-10), left posterior (6-8-11-13-9); right anterior (6-9-12-11- 11), left anterior (6-10-13-11-12), fourth finger of the extremities longer than others; digital toepads longer than wide on all fingers. Measurements in mm: snout-vent length 70.9; axilla-groin 35.8; head length 16.5; head width 13.2; snout length (to eye) 10.05; eye diameter 4.1; auricular opening (maximum) 1.6; length of fourth finger 5.5; length of fourth toe 5.7; width between eye supercilliaries 9; internaral 2.1. Coloration of holotype in ethanol (Fig. 8). The holotype exhibits relatively little coloration, the dorsal background color varies from brown to creamy yellow, with dark brown faint irregular spots (less evident compared to live coloration), the ventral surface varies yellowish and cream; half of the ventral color on the tail to the tip is cream. Pattern and color in life (Fig. 10). Dorsally presents a clear background, a mixture of cream-pink, on this background, patterns of irregular patches dark brown color and some completely black spots are present, some of these spots are slightly surrounded by yellowish scales; on the middle dorsal part a longitudinal line of the background color runs from head to tail; scales around the eyes and some supralabials are slightly colored light yellow; ventrally presents a white-cream coloration in combination with light yellow, more evident towards the ventral region; the pupils are black on a bronze iris. Dorsal coloration of the tail is white-cream with brown stripes and some black points. Variation. All meristic and morphometric characters are presented with mean values, standard deviation and range of meristic (Table 2) and morphometric variables (Table 7). Distribution and habitat. Phyllodactylus benedettii sp. nov. is an endemic species to the state of Jalisco, restricted to the Chamela-Cuixmala Biosphere Reserve. According to the INEGI digital map of elevation (resolution of 15 m), the type locality is located at 100 masl, located in the biogeographic province of the Western Pacific surrounded by deciduous tropical forests (Fig. 11). The microhabitats reported for this species are diverse: soil, trees, shrubs and in habitats modified by man as houses or bridges or some other constructions (meteorological stations). They are particularly abundant in rocks and crevices and in people's houses or buildings where it feeds on insects and other invertebrates (García & Ceballos 1994; Ponce-Campos & García 2007; Vitt & Caldwell 2014; pers. obs.). Males of P. benedettii have an extended period of reproduction (August–March) with two peaks, one in rainy season (August–October) and another one in dry season (November–March). Reproductive behavior (courtship and mating) begins in August which coincides with the beginning of rainy season. The females produce clutches of two eggs, with up to three clutches during the breeding season, with a peak of egg production from December to March coinciding with the dry season (Ramírez-Sandoval et al. 2006). This species presents a distinct karyotype from P. lanei presumably due to Robertsonian fusions/fissions with 19 pairs of telocentric chromosomes (2n = 38, FN = 38) (Castiglia et al. 2009). They are lizards of nocturnal habits and feed on insects, although to date there have been no studies on the composition of their diet (García & Ceballos 1994; Vitt & Caldwell 2014). Representative species of tropical dry forest of Chamela-Cuixmala Biosphere Reserve are: Sciadodendrom excelsum, Brosimum alicastrum, Orbignya cohune and Tabebuia donell-smithi (Ceballos et al. 1999). The deciduous tropical forests is near the mouth of the Cuixmala River and is characterized by the abundance of water in the rainy season, the climate is tropical (warm-humid), has an annual average temperature of 25 °C and the annual rainfall varies from 748–1000 mm (García & Ceballos 1994; Ceballos et al. 1999). Other species of reptiles with nocturnal activity that inhabit the deciduous tropical forests are: Hemidactylus frenatus, Boa constrictor, Lampropeltis triangulum, Leptodeira maculata, Pseudoficimia frontalis, Senticolis triaspis, Tropidodipsas philippii (Sibon philippi), Tantilla bocourti, Thamnophis valida, Trimorphodon biscutatus, Micrurus distans, Agkistrodon bilineatus, and Crotalus basiliscus (García & Ceballos 1994). Comments on conservation and threats. This species inhabits the Chamela-Cuixmala Biosphere Reserve decreed in 1993. There is a minor threat posed by the localities bordering the reserve, and according to the observations made by the first author, apparently there is a large population of P. benedettii geckos inside Chamela- Cuixmala Biology Station (UNAM). There are two main threats: the false beliefs of some people who consider this species as poisonous (local inhabitants), and the possible negative effects due to the presence of invasive gecko Hemidactylus frenatus, a species with a very large population that may exploit the same ecological niche. To date there is no specific study on niche overlap between H. frenatus and Phyllodactylus species in Mexico, however, its potential danger to local biodiversity has been documented, including interspecific negative interactions with native geckos from different areas of the world; one example of this is the displacement and reduction of six gecko species of the genus Nactus on the Mascarene Islands and Lepidodactylus lugubris throughout Pacific islands. In these cases H. frenatus has been observed stalking, lunging towards and biting at other geckos (Global Invasive Species Database 2015). Due to the above, and as a potential risk to local biodiversity in Mexico, it is classified as a high risk invasive species (CONABIO 2016). Phyllodactylus kroPotkini sp. nov. Holotype. Adult male (MZFC 28736) collected in Nueva Filadelfia (Huerta Vieja) in the municipality of Tlapehuala, Guerrero (18.29 N, - 100.49 W WGS84), collected on March 15, 2014 by Tonatiuh Ramírez Reyes and Joel Rosas Avila. Paratypes (4). Four male adults (MZFC 28735, MZFC 28737, MZFC 28738, MZFC 28739) collected at the type locality, Nueva Filadelfia (Huerta Vieja), Tlapehuala, Guerrero (18.29 N, - 100.49 W WGS84) collected on March 16, 2014 by Tonatiuh Ramírez Reyes and Joel Rosas Ávila. Etymology. The species is dedicated to the memory of the great Russian philosopher, geographer and naturalist Piotr Kropotkin, who made great scientific and theoretical contributions about mutual support and altruism in some animal populations (including human society). Diagnosis. Phyllodactylus kropotkini sp. nov. is a medium-sized gecko within the P. lanei complex. Dixon (1964) established three fundamental characteristics to include species and subspecies within the P. lanei complex and differentiate them from P. tuberculosus, species of the P. lanei complex have a low number of interorbital scales (12–19), white venter and low number of scales across the snout between the third labials (does not specify range of values). Phyllodactylus kropotkini has 14.6 interorbital scales on average (14–15), white venter and 20.2 third labial–snout scales on average (19–21). Phyllodactylus kropotkini presents a maximum recorded SVL of 62.3 mm that clearly differs from the nearest species P. lanei (max. 71 mm) and it differs from other species: P. isabelae (max. 57.5), P. lupitae (max. 73.3 mm), P. rupinus (max. 69.4), P. benedettii (max. 74.2) and P. bordai (max. 58.9 mm). Phyllodactylus kropotkini sp. nov. shows the following combination of characters: 28 (25–31) tubercles from head to tail, 67 (63–73) longitudinal scales, 13 (12–14) rows of tubercles across dorsum, 30 (29–33) scales across venter, 14 (14–15) interorbital scales and 20 (19–21) third labial–snout scales. Phyllodactylus kropotkini has a partial overlap with P. lanei and P. isabelae on morphometric space (Fig. 7), however P. kropotkini is clearly differentiated from P. lanei by maximum length (SVL) (Fig. 6); it also presents the follow measures: axilla-groin length (22.54 mm), snout length (9.34 mm), auricular opening (1.78 mm), while P. lanei has 24.7 mm axilla-groin length, 10. 61 mm snout length and auricular opening (1.95 mm) and P. isabelae has 18.7 mm axilla-groin length, 8.18 mm snout length and auricular opening 1.51 mm. P. kropotkini has a low number of interorbital scales (14–15, 14.6 avg.) similar to P. benedettii (13–16, 14.4 avg.), all other species of Phyllodactylus exceed 15 interorbital scales on average, P. isabelae (14–16, 15.1 avg.), P. lupitae (14–20, 16.83 avg.), P. lanei (15–17, 15.5 avg.), P. rupinus (14–17, 15.5 avg.), P. t. magnus (21–23, 22.3 avg.), P. muralis (21–27, 23.2 avg.), P. tuberculosus (16–22, 19.2 avg.). There are an average of 30.6 scales across the venter (29–33), unlike P. isabelae (26–29, 27.8 avg.), P. lupitae (23–26, 24.6 avg.), P. lanei (29–32, 30 avg.), P. rupinus (24–28, 26.5 avg.) and P. benedettii (25–32, 29.1 avg.). Phyllodactylus kropotkini presents 67.6 longitudinal scales from the gular region to the anus on average (63–73), compared to P. rupinus (63–73, 63.6 avg.), P. lanei (60–74, 66 avg.), P. benedettii (60–69, 62.6 avg.), P. t. magnus (52–58, 54.6 avg.), P. mura

Crotalus thalassoporus sp. nov. (Figure 5; Table 3) Crotalus mitchelli pyrrhus, Seib 1978, in part. Crotalus mitchellii, Grismer 2002b, Murphy and Aguirre-Léon 2002, Meik et al. 2012a, in part. Crotalus pyrrhus, Meik et al. 2015, in part. Description A diminutive insular species of speckled rattlesnake with overall pale appearance; background colouration tan to pinkish with a series of 27–40 indistinct blotches; blotches usually only slightly darker than ground colour, pinkish to pale brown, often appearing faintly rust-coloured; dark speckling on body is faint. Tail with 3–5 bands, last two or three distinctly black with pale cream interspaces. The head is similar in colouration to the body, but often with few dark specks, more conspicuous than stippling on trunk; grey postocular stripe indistinct. In colour pattern, C. thalassoporus is similar to C. angelensis, and in both species the anterior scales of the dorsal body blotches are tipped posteriorly with dark brown or black. A single row of nasorostral scales precludes contact between rostral and prenasal scales. Variation in standard phenotypic characters is presented in Table 3. Diagnosis The presence of nasorostral scales distinguishes the new species from all congeners except for species of the C. mitchellii complex. From C. mitchellii the new species differs in having typically fewer dorsal scale rows (21.5–22 vs 25), fewer subcaudals (16–22 vs 20–24), fewer temporal scale rows (7 vs 8), fewer supralabials (14 vs 16), fewer interrictals (22–23 vs 28), fewer prefrontals (18–20 vs 30–31), colour pattern (pale tan, pinkish, or beige ground colour with indistinct rust-brown blotches vs variable colour pattern), and smaller adult body size. From mainland populations of C. pyrrhus the new species differs in having typically fewer tail bands (3–4 vs 4–6), fewer dorsal scale rows (21.5–22 vs 25), fewer temporal scale rows (7 vs 8), fewer supralabials (14 vs 16), fewer interrictals (22–23 vs 29), fewer prefrontals (18–20 vs 27–34), colour pattern (pale tan, pinkish, or beige ground colour with indistinct rust-brown blotches vs extremely variable), and smaller adult body size. From C. angelensis, the new species differs in having typically fewer tail bands (3–4 vs 5–8), fewer dorsal body blotches (31–32 vs 40–41), fewer dorsal scale rows (21.5–22 vs 27), fewer subcaudals (16–22 vs 20–25), fewer ventrals (169.5–174 vs 182–188), more supralabials (14 vs 13), fewer interrictals (22–23 vs 29–30), fewer prefrontals (18–20 vs 25–26), and smaller adult body size. From C. polisi, the new species differs in having typically fewer tail bands (3–4 vs 5–6), fewer dorsal body blotches (31– 32 vs 40–42.5), fewer subcaudals in females (16 vs 17), more ventrals in females (174 vs 169), fewer interrictals in males (22 vs 26), fewer prefrontals (18–20 vs 21–27.5), and colour pattern (pale tan, pinkish or beige ground colour with indistinct rust-brown blotches vs slate or charcoal grey ground colour with indistinct blotches). Holotype Subadult male, MZFC-26410, field number JMM-648, collected on 19 March 2010 by Jesse M. Meik, Sarah Schaack and Matthew J. Ingrasci. Rostral plate is as high as broad (2.2 × 2.2 mm), separated from the prenasal scales by 3/2 nasorostral scales; internasal scales 2, with an additional tiny scale interpositioned at the anterior suture and in contact with dorsal edge of rostral. Head scalation highly irregular, making some scale designations ambiguous. Distinct canthal scales absent, but approximately 22 knobby scales of variable size and shape in prefrontal area; interocular scales 7; loreal scales 3/3, irregularly shaped; preocular scales 2/2, prefoveal scales 11/8, irregularly shaped, precluding contact between nasal scales and supraocular scales; subocular scales separated from supralabials by two scales at midpoint of eye; supralabial scales 15/15; infralabial scales 17/16; interrictal scales 24; dorsal scale rows at midbody 23; ventrals 169 (exclusive of three preventrals); subcaudal scales 21, undivided (except distal 3, which are divided); rattle fringe scales 10; rattle segments 3, chain incomplete. Measurements. SVL, 341 mm; tail length, 28 mm; head length (rostral plate to articulation of mandible with quadrate), 18.5 mm; head width (at widest point just anterior to articulation with mandible), 15.7 mm; proximal rattle segment width, 6.7 mm. Colouration and pattern in preservative. Overall ground colouration dirty cream with faint speckling; head with few faint grey specks; lateral surfaces of head with medium grey suffusion, labial scales with cream spots; ventral surface of head cream, immaculate, ventral surface of trunk cream with diffuse black specks; 37 indistinct body blotches only slightly darker than ground colour, some with pale centres, all primary dorsal blotches wider than long with exception of first four, fusing with faint lateral blotch series over posterior third of body to form crossbands; dark maculations on anterior and posterior margins of blotches give the impression of faint transverse bars along the length of the body; five tail bands, distal three black and two scales long. Type locality Piojo Island, Municipality de Ensenada, Baja California, Mexico. Coordinates: N 29.018 W 113.465 (Figure 5). Type deposition Holotype at MZFC-UNAM; paratypes at MZFC-UNAM (MZFC 26411, MZFC 26412) and at UTAARDRC (UTA R-59766, UTA R-59767). Etymology The specific name is derived from the Greek word meaning ‘seafarer’, and is a reference to the apparent historical introgression we note between this taxon and the population of speckled rattlesnakes on Smith Island, most likely resulting from oversea dispersal of propagules from Piojo Island (see Discussion).

Crotalus polisi sp. nov. (Figure 5; Table 3) Crotalus mitchellii, Grismer 2002a, Murphy and Aguirre-Léon 2002, Meik et al. 2012a, in part. Crotalus pyrrhus, Meik et al. 2015, in part. Description A diminutive insular species of speckled rattlesnake with overall dusky appearance; background colouration is medium grey with a series of 36–48 indistinct and irregularly shaped dorsal body blotches, usually slate to charcoal grey in colour, only slightly darker than background; tail with 5–8 bands, last several are black with cream interspaces. The head is similar in colour to the body, with dark lateral suffusion and often with a faint postocular stripe and parietal blotches. Pattern is heavily punctated with black specks; blotches are usually wider than long, and merge with secondary lateral series to form muted crossbands on the posterior half of the body. A single row of nasorostral scales precludes contact between rostral and prenasal scales. Variation in selected phenotypic characters is presented in Table 3. Diagnosis The presence of nasorostral scales distinguishes the new species from all congeners except for species of the C. mitchellii complex. From C. mitchellii the new species differs in typically having more tail bands (range of mode or median between males and females of each species is presented for all comparisons: 5–6 vs 3–4), more dorsal body blotches (40–42.5 vs 32–34), fewer dorsal scale rows (23 vs 25), shorter ultimate supralabial scale (slightly longer than high vs twice as long as high), fewer supralabials (14 vs 16), fewer ventral scales in females (169 vs 180), fewer temporal scale rows (7 vs 8), colour pattern (mostly uniform colour pattern of slate or charcoal grey with indistinct blotches vs variable colour pattern), and smaller adult body size. From mainland populations of C. pyrrhus the new species differs in having typically more dorsal body blotches (40–42.5 vs 33–34), fewer dorsal scale rows (23 vs 25), fewer ventrals (168–169 vs 176), fewer temporal scale rows (7 vs 8), fewer supralabials (14 vs 16), colour pattern (mostly uniform colour pattern of slate or charcoal grey with indistinct blotches vs extremely variable), and smaller adult body size. From C. angelensis the new species differs in having typically fewer dorsal scale rows (23 vs 27), more supralabials (14 vs 13), fewer ventrals (168–169 vs 182–188), colour pattern (colour pattern of slate or charcoal grey with indistinct blotches vs buff or pink ground colour with grey to russet hexagonal blotches), and smaller adult body size. From C. thalassoporus, the new species differs in having more tail bands (5–6 vs 3–4), more dorsal body blotches (40–42.5 vs 31–32), more interrictals (25–26 vs 22–23), more prefrontals (21–27.5 vs 18–20), and colour pattern (colour pattern of slate or charcoal grey with indistinct blotches vs fawn, pinkish or beige ground colour with indistinct rust-brown blotches). Holotype Adult female, MZFC-26408, field number JMM-642, collected on 18 March 2010 by Jesse M. Meik, Sarah Schaack and Matthew J. Ingrasci. Rostral plate slightly broader than high (2.5 × 2.2 mm). Head scalation highly irregular, making some scale designations ambiguous. Rostral-prenasal contact precluded by 3/3 nasorostral scales; two internasals contact rostral; distinct canthal scales absent, but approximately 29 knobby scales of variable size and shape in prefrontal area; interocular distance spans a minimum of six scales; loreal scales 4/4, irregularly shaped; preocular scales 2/2, upper prefoveal scales irregular, lower prefoveal scales large and broadly contact first three supralabial scales on both sides; subocular scales separated from supralabials by three scales at midpoint of eye; supralabial scales 14/14; infralabial scales 15/15; interrictal scales 25; dorsal scale rows at midbody 23; ventrals 168 (exclusive of one preventral); subcaudal scales 17, undivided; rattle fringe scales 12; rattle segments 6, button present. Measurements. Snout–vent length (SVL), 445 mm; tail length, 23 mm; head length (rostral plate to articulation of mandible with quadrate), 20.7 mm; head width (at widest point just anterior to articulation with mandible), 19.9 mm; proximal rattle segment width, 8.5 mm. Colouration and pattern in preservative. Head with diffuse black speckling; lateral surfaces of head dusky with faint postocular stripe; labial scales with cream blotches (appearing as ‘blotch negatives’ against surrounding ground colour); ventral surface of head cream with faint black specks on periphery, ventral surface of body also cream with diffuse black specks, becoming more prominent on posterior one fourth of body; 38 dusky grey body blotches with narrow (0.5–2 scales long) cream interspaces; blotches indistinct with ill-defined borders, all wider than long, fusing with faint lateral blotch series over posterior third of body to form crossbands; five tail bands, distal four black and only one to two scales long. Type locality Cabeza de Caballo Island, Municipality de Ensenada, Baja California, Mexico. Coordinates: N 28.971 W 113.479 (Figure 5). Type deposition Holotype at MZFC-UNAM; paratypes at MZFC-UNAM (MZFC 26407, MZFC 26409) and at UTAARDRC (UTA R-59763, UTA R-59764, UTA R-59765). Etymology The specific name is a patronym honouring the late Gary A. Polis of the University of California Davis, a renowned arachnologist and desert food-web ecologist, who died at sea on 27 March 2000 when his research vessel capsised in a gale while returning to Bahía de Los Angeles from an expedition to Cabeza de Caballo Island. In addition to Polis, four other researchers, including postdoctoral fellow Michael D. Rose of UC Davis, and Takuya Abe, Masahiko Higashi and Shigero Nakano of Kyoto University, Japan, perished on that day. Four other UC Davis researchers and students survived the tragedy, and by their accounts, the deceased heroically gave their lives to help ensure their survival., Published as part of Meik, Jesse M., Schaack, Sarah, Flores-Villela, Oscar & Streicher, Jeffrey W., 2018, Integrative taxonomy at the nexus of population divergence and speciation in insular speckled rattlesnakes, pp. 989-1016 in Journal of Natural History 52 (13 - 16) on pages 1003-1005, DOI: 10.1080/00222933.2018.1429689, http://zenodo.org/record/5174557, {"references":["Grismer LL. 2002 a. A re-evaluation of the evidence for a mid-Pleistocene mid-peninsular seaway in Baja California: a reply to Riddle et al. Herpetol Rev. 33: 15 - 16.","Murphy RW, Aguirre-Leon G. 2002. Distributional checklist of nonavian reptiles and amphibians on the islands in the Sea of Cortes. In: Case TJ, Cody ML, Ezcurra E, editors. A new island biogeography of the Sea of Cortes. New York (NY): Oxford University Press; p. 580 - 594.","Meik JM, Schaack S, Ingrasci MJ, Lawing AM, Setser K, Mocino-Deloya E, Flores-Villela O. 2012 a. Notes on activity, body size variation, and diet in insular speckled rattlesnakes from the western Sea of Cortes, Mexico. Herpetol Rev. 43: 556 - 560.","Meik JM, Streicher JW, Lawing AM, Flores-Villela O, Fujita MK. 2015. Limitations of climatic data for inferring species boundaries: insights from speckled rattlesnakes. PLoS One. 10: e 0131435."]}

Hylodes augusti Dugès, 1879 a The species was originally described by Dugès (1879 a) based on an abbreviated version of a manuscript he wrote in 1868, the latter being published in its entirely by Brocchi (1881). The original description of 1879 makes reference to only one specimen from Guanajuato, Mexico (Dugès 1879 a: 21–22). Brocchi’s (1881) expanded version of Dugès’ manuscript of August 1868, was based apparently on two specimens, although since there was only one specimen noted in the original description of 1879 a, the second specimen has no status as a type. Additionally, there are measurements for one specimen and illustrations in plate 16 figures 1, 1 a– 1 d of Brocchi (1881). It is important to note that plate 16 was not published until 1883 (Smith & Smith 1973). Smith & Necker (1943: 201) stated that “…The original description was apparently based upon three cotypes…” (Dugès 1879 a). We see no evidence that this was the case. At most, there are two specimens in Dugès’ description in Brocchi (1881): one specimen from Guanajuato, presumably the holotype noted in the original description, and a second from Tupátaro from a hole at the end of a cave called the “Cueva del Padre Torres” in the middle of Cerro San Gregorio Hill (Cueva del Padre Torres, au milieu du Cerro San Gregorio), Guanajuato. According to Smith & Necker (1943), none of the cotypes (= syntypes) were present at the MADUG when they examined the collection. Although they found one specimen that had a tag marked by Dugès as “ type ”, it was collected at a depth of 40 m in the Cedro Mine, Guanajuato; the specimen has no original tag but a typed tag (inside the jar) has the verbatim information reported by Smith & Necker (1943: 201). They designated this female specimen as a neotype and gave a detailed description of it. This specimen is catalogued as HE 324 (Fig. 1 A). Smith & Necker (1943) designated it as a neotype because they were unable to find any of the syntypes. Later, Zweifel (1967) noted that the type was a skeleton following Smith & Taylor (1948); this information was originally reported by Mocquard (1899: 161) who stated “J’avais en outre appris depuis peu que M. Dugès, entre les mains duquel était resté le type de H. Augusti, avait eu la malencontreuse idée de le sacrifier pour préparer le squelette! Je désespérais du pouvoir résoudre la question, lorsque l’envoi tout récent, fait au Muséum par ce naturaliste, d’un jeune spécimen d’ H. Augusti, vint heureusement me tirer d’embarras” (I recently learned that Mr. Dugès, who had in his hands the type of H. Augusti, had the unfortunate idea of sacrificing it to prepare the skeleton! I despaired of ever resolving the matter, when this naturalist recently sent a juvenile specimen of H. Augusti to the Museum, which happily helped me answer the question). There are two skeletons at the MADUG (HE 999, 1000; Fig. 1 B–C). Based on the measurements given in Brocchi (1881) HE 1000 is the specimen that most closely matches those measurements. It is important to mention that the original tag is not found with any of these specimens, only a note, not in Dugès’ handwriting, that says Moroleón, Gto. We believe that HE 1000 is the original type (Fig. 1 C). Therefore, the neotype (HE 324) was wrongly designated by Smith & Necker (1943). Current valid name: Caugastor a. augusti (Dugès in Brocchi, 1879)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on pages 34-35, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Crotalus jimenezii Dugès, 1877 In the original description of this taxon, Dugès (1876–1877) referred to more than one specimen, although he only gave measurements for one individual. There are two surviving specimens at the MADUG (HE 55, 356; Figs 8 D, 9 A). The latter is the same one reported by Smith & Necker (1943). We think these specimens should be considered syntypes together with those from the US National Museum (USNM 24448, 26152 and 46508), designated by Smith & Necker (1943). Both specimens were given to Dugès by Dr. Villalobos and have original tags with Dugès handwriting; HE 55 (Fig. 8 D) is from Guanajuato, and HE 356 (Fig. 9 A), from “S. Ant. el Rico” (San Antonio el Rico, Guanajuato). There are also two additional skulls, HE 1141, 1142 (Fig. 9 B–C) that also should be considered syntypes since Dugès (1876–1877) stated “Los palato-maxilares son de una cuarta parte más largos que la porción del terigoides comprendida entre la extremidad posterior del palato-maxilar y la articulación del terigoides con el hueso timpánico.” (The palato-maxillaries are one quarter longer than the portion of the pterygoid between the rear end of the palato-maxillary and pterygoid articulation with the tympanic bone.) Our measurements of this character are: HE 1141 (Fig. 9 B) palato-maxillary = 15.7 mm, last portion of pterygoid = 12.7 mm; HE 1142 (Fig. 9 C) palatomaxillary = 14.5 mm, last portion of pterygoid = 11.4 mm. There are four additional specimens from “Guanajuato” that have been considered syntypes; USNM 24448 and USNM 26152, mentioned by Smith & Necker (1943) and MNHN 1883.284 and MNHN 1883.288, mentioned by Thireau (1991). The first two were not mentioned in the type catalogue of the Smithsonian by Cochran (1961) and the other two have a question mark as syntypes by Thireau (1991). All these specimens could be syntypes, although they do not have a more specific locality; Dugès mentioned in his description of this taxon, “He recibido este ofidio de Silao, Colima y Guadalajara…” (I have received this ophidian from Silao, Colima and Guadalajara…). It is clear that Dugès used a series of specimens in his description of this taxon, some of them may have been exchanged with other museums, like the Smithsonian and the Muséum National d’Histoire Naturelle in Paris. Finally, regarding the publication date for this name, McDiarmid et al. (1999) discussed the correct date for the publication of this new taxon by Dugès. Since this paper was published over two years in different “entregas”, pages 23–25, were the description of Crotalus jimnezi appeared was part of “entregas” 3 and 4, but the name was published on page 23, belonging to 1877. Current valid name: Crotalus polystictus (Cope, 1865)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 44, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Oreophis boulengeri Dugès, 1897 The holotype is still at the MADUG and was described by Smith & Necker (1943). This specimen has the number HE 103 (Fig. 8 A). Current valid name: Lampropeltis mexicana (Garman, 1884)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 43, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Siredon dumerilii Dugès, 1870 We found only one specimen in MADUG (HE 315; Fig. 4 C). When Smith & Necker (1943) reported on the type collection they did not find this specimen, which was subsequently reported on by Maldonado Koerdell (1948 a, b). Two additional specimens reported by Smith & Necker (1943) had the locality San Juan del Río, Guanajuato (see comment under Bathysiredon dumerilii queretarensis). Maldonado Koerdell (1948 b) corrected the locality to San Juan del Río, Querétaro, and described these two specimens as Bathysiredon dumerilii queretarensis (see below). Dunn (1939), mentioned three additional specimens that are considered cotypes (syntypes): ANSP 13862, presented by Dugès to Cope; and USNM 16201 – 2. None of these was specifically designated by Dugès (1870, 1872) as a type specimen. It is obvious that the description was based on at least one female and one male, but considering the number of specimens now recognized as syntypes, Dugès may have had more than two specimens when he described the species. There is an additional specimen of this taxon incorrectly labeled as coming from Colima, Mexico (HE 546, Fig. 4 D); the tag is not original, so we conclude it may have been erroneously labeled at a later date. In order to avoid any confusion, we do not assign any type status to this particular specimen. Examination of the specimen HE 315, mostly coincides with Maldonado Koerdell’s (1948 a) description, although the measurements are not exactly the same as those he reported. Our measurements, with Maldonado Koerdell’s in parenthesis, are: total length = 264mm (275), length of head and trunk = 135.5 (140), length of left anterior limb = 47 (45), length of left posterior limb = 46 (51). Also, one of us (OFV) counted 13 costal grooves instead of the 14 reported by Maldonado Koerdell (1948 a). Current valid name: Ambystoma dumerilii (Dugès, 1870)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 38, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Hemigenius variabilis Dugès, 1888 d The species and the genus were apparently described on the basis of one specimen. One specimen, with no locality data according to Smith & Necker (1943) was present at MADUG when they reviewed the types of this collection. They redescribed it and presented photographs of the head and neck. Nevertheless, Dugès (1888 d) mentioned “J’ai trouvé ce petit serpent à la Barranca de Portillo (Guadalajara) et aussi à Guanajuato.” (I found this little snake at the Barranca of Portillo (Guadalajara) and also in Guanajuato); this statement leads us to believe that Dugès had more than one specimen at hand when describing the species. We were unable to find the specimen described by Smith & Necker (1943), despite checking all possible candidates in the collection; none coincides with measurements given by Smith & Necker (1943). The type is definitively lost. Current valid name: Storeria storerioides (Cope, 1865)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on pages 47-48, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Adelophis copei Dugès 1879 b The original description for this taxon appeared in Cope (1879); Cope credited the description to Dugès in manuscript. Although Cope did not reproduce a verbatim translation of Dugès’ manuscript, published later by Dugès himself (1887), he did copy all of the measurements and scale counts as they are reported in Dugès (1887). In brief, Dugès’ (1887) description has a little more text than Cope’s translation. Additionally Duges’ justification for publishing the description in “La Naturaleza” was that “El sabio herpetólogo E. D. Cope…ha traducido una nota que le transmití sobre este ofidio, pero sin acompañarla con la debida lámina; por este motivo me ha parecido conveniente publicarla con las figuras necesarias, para poder reconocer á primera vista este reptil, raro…” (The wise herpetologist E. D. Cope…has translated a note that I conveyed him about this ophidian, but he did not accompany it with the proper plate; for this reason it is proper for me to publish it (referring to the description) with the necessary figures, in order that one might to be able to recognize this rare reptile at first sight …). The type locality was stated in both publications as Guadalajara. A caveat to this matter is that Dugès’ (1887) description, was dated as “Guanajuato, Octubre 22 de 1880 ”, and Cope’s (1879) appeared in the June 20 issue of the Proceedings of the American Philosophical Society. According to the requirements of the ICZN (1999) Article 5.1. 1 regarding availability of the source, it is clear that Cope did not have a specimen available on which to base his description as there are no candidate type specimens of Adelophis at the ANSP (Malnate 1971) nor at the USNM (Cochran 1961); rather, he probably based it on a preliminary draft obtained from Dugès. When Smith & Necker (1943) reviewed the types at MADUG, they found one specimen of this taxon, from “Cupátaro”, probably a typo for Tupátaro, Guanajuato; they decided to designate it a neotype, because Dugès (1887: 20) stated that he did not have the specimen in his custody, and did not mention the whereabouts of the specimen used by him to describe this taxon. In that same publication (Dugès 1887), is also a mention of a well preserved skull of this species, that Dugès had in his possession; this skull is labeled HE 0 963 (Fig. 10 D). This specimen cannot be considered the type since there is no drawing of the skull in the publication, only a mention of its existence in the re-description of the taxon by Dugès (1887) in “La Naturaleza”. Unfortunately, there is no locality information accompanying this skull. Specimen HE 0 963 is of value since it is the only surviving specimen of this rare taxon in the MADUG. The type is the specimen, now lost or destroyed, figured in Plate III fig 1 a-d in Dugès (1887) as this was the specimen used for the description published in Cope (1879) and Dugès (1887), and originating from the type locality of Guadalajara. Current valid name: Adelophis copei Dugès in Cope, 1879, Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 47, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Bathysiredon dumerilii queretarensis Maldonado Koerdell, 1948 Maldonado Koerdell (1948 b) reported two specimens that Smith & Necker (1943) described as Ambystoma larvae. These specimens were labeled from “San Juan del Río, Guanajuato”, according to Smith & Necker (1943). But Maldonado Koerdell (1948 b) stated that they were from “San Juan del Río (Qro.)”, arguing that Smith & Necker (1943) put them in the state of Guanajuato rather than in Queretaro, where a city called San Juan del Río is well known. Although there is a small town named San Juan del Río in the state of Guanajuato, in the municipality of Pénjamo (N 20.314029º, W 101.734933º, INEGI 2015) we do not believe that the specimens could have come from this locality, because there is no a lake there (only a dam, probably recently built) and the population of this town is only 38 (http://www.microrregiones.gob.mx/catloc/contenido.aspx?refnac= 110230632). Maldonado Koerdell referred to these specimens as cotypes (syntypes) and described them; they are catalogued as HE 1184–1185 (Fig. 9 D, 10 A). Brandon (1992) questioned the validity of this taxon and provides ample discussion on the matter. We concur with Brandon (1992) that this taxon should be synonymized with A. dumerilii. However, the specimens shown in Figures 9 D, 10 A do not coincide with Maldonado-Koerdell’s description; one of us (OFV, personal notes) checked the holotypes in 1985 and his recollection is that they looked like A. dumerilii. We do not believe that specimens HE 1184–1185 (Fig. 9 D, 10 A) correspond to the taxon in question. Our conclusion is that both syntypes are lost. Current valid name: Ambystoma dumerilii (Dugès, 1870)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 45, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Liophis janii Dugès, 1865 The type specimen had apparently been lost by the time Smith & Necker visited the collection, and they designated a topotype from some material that they had previously examined in the US National Museum (USNM 9889) as the neotype (Smith & Necker 1943). They did not describe the neotype, but rather transcribed the original description in French by Dugès. Tanner (1946: 49) also commented that two specimens in the USNM (9889 and 11369) had been collected by Dugès from Guanajuato, Mexico and “ … may represent type material ”, thereby independently reaching the same conclusion as did Smith and Necker. There is no indication in Dugès description (1865) that he had sent any other specimen to the USNM or ANSP (Cochran 1961; Malnate 1971). According to Dugès (1865: 32), the specimen was from Guanajuato, and a comment at the end of the paper “Quelquefois la tache du cou est séparée en trois, deux latérals et la languette médiane” (Sometimes the neck spot is separated in three parts, two at the sides and the stripe in the middle) suggests that he had more than one specimen when describing this taxon. We found a specimen at MADUG (HE 102, Fig. 7 C) from Guanajuato that has more or less the same dimensions as those given by Dugès in the original description (head width: 10.2 mm, head length: 14.5mm, SVL: 300mm, body width: 14mm, tail: 53mm); this specimen also has the characteristic neck marks as described by Dugès (1865: 32). The original tag bears the locality Guanajuato, as in the original description. We consider this specimen to be the holotype and the one that Dugès originally described. Therefore, the designation of a neotype by Smith and Necker was incorrect. Current valid name: Hypsiglena janii (Dugès, 1865)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 42, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Amblystoma altamirani Dugès, 1895 This is one of the taxa described by Dugès from more than one specimen. The original work was published as part of a report for the “Secretaría de Fomento” (Ministry of Development) by Dr. Fernando Altamirano (1895). The original description was first published in French (pp. 5–9) and later translated into Spanish (pp. 9–13). Altamirano indicated in the French description that he had about 20 specimens, but in the Spanish section he mentioned more than 20. In the Introduction to Zoology section of the report, Altamirano stated that he had sent about 25 specimens to Dugès. Smith & Necker (1943) found six specimens preserved in alcohol; the same number that can be found today at the MADUG (HE 314, 316–319, 322; Figs 1 D, 2 A–D, 3 C). Besides these specimens, there are two others (Fig. 3 A–B), a skull and appendages (HE 964) and a skull (HE 965). These specimens were not reported by Smith & Necker (1943) but represent part of the figures in the original description. Therefore, specimens HE 964 and HE 965 should be considered members of the syntype series. Smith & Necker (1943) also redescribed a specimen they considered to be most similar to the illustration in the original description, this can be either HE 316 or HE 319, both are very similar and are completely metamorphosed as is that illustrated by Dugès. Current valid name: Ambystoma altamirani Dugès, 1895., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on pages 36-37, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Rhadinella dysmica sp. nov. Figs. 1���2, Table 1 Holotype. Adult female, ENCB 18951, obtained 17 July 2014 by L. F. D��vila Galav��z near cueva de Tepozonales (17.2853 N, ��� 99.3662 W; 432 m above sea level, asl hereafter), Cuajilotla, 20 km S Mochitl��n, Guerrero, Mexico. The individual was found near the entrance of the cave, crawling on the ground. Diagnosis. A member of Rhadinella, genus characterized by gracile body, head only slightly distinct from neck; inverted dark U-shaped marking on rostral; anterior supralabials usually with dark edges and pale centers; conspicuous or at least evidence of pale collar on neck; with reduced to small supracloacal tubercles in adult females; no subpreocular scale; posterior (2���7) teeth larger and thicker than preceding teeth, all enlarged teeth on same plane, no broad diastema; with basal nude pocket on hemipenis. Rhadinella dysmica is further characterized by the combination of a distinctive white collar, undivided at middorsum, extending 2.0��� 2.5 scale lengths at middorsum and 3���4 scale lengths on side of head; temporals 1 + 2; smooth dorsal scale rows 17, unreduced throughout body; ventrals 172; cloacal scute divided; divided subcaudals incomplete 63 +N; dorsal coloration dark brown to blackish, almost totally obfuscating longitudinal striping, ground color barely extending to lateral edges of ventrals. Rhadinella dysmica may be distinguished from many congeners (i.e., R. godmani, R. hannsteini, R. hempsteadae, R. kanalchuchan, R. kinkelini, R. lachrymans, R. montecristoi, R. pegosalyta, R. rogerromani, R. serperaster, and R. tolpanorum) by dark dorsal body coloration and absence of conspicuous striping. Additionally, the new species differs from R. godmani, R. hempsteadae, R. kanalchuchan, R. montecristoi, R. pegosalyta, and R. serperaster by having 17 dorsal scale rows (vs. more than 17 dorsal scale rows). Rhadinella dysmica is morphologically similar to other species having mostly uniformly blackish dorsum, with or without indistinct dark vertebral line and pale dashes on some dorsal scales (i.e., R. schistosa, R. donaji, R. pilonaorum, and R. posadasi). In R. dysmica the color and configuration of the collar alone diagnostic from all other species of dark Rhadinella, extending from about posterior tips of parietals for several scale lengths, being white, and distinctly set off from mostly black head-cap. In R. shistosa collar U-shaped and usually broken medially (Fig. 3); in R. donaji white coloration of collar extends on to head, involving most of parietals and part of frontal; in R. pilonaorum collar orange with top of head mostly auburn mottled with black; in R. posadasi the collar is dark orange or auburn. Other features distinguishing these dark-colored species of Rhadinella include fewer ventrals (151���156 in females), fewer subcaudals (31���34 in females), and 1 + 1 temporals in Rhadinella shistosa; more subcaudals (100���111 in females) in R. pilonaorum; and fewer ventrals (143���146 in females) and more subcaudals (86���93 in females) in R. posadasi. Fourth and fifth infralabials contact posterior pair of genials in R. schistosa, R. pilonaorum, and R. posadasi (vs. the third and fourth in R. dysmica and R. donaji); distal portion of tails incomplete in holotypes of R. dysmica (a female) and R. donaji (a male), however with 66 subcaudals R. dysmica appears to have higher number than R. donaji, which has 40. Description of the holotype. Adult female with few small supracloacal tubercles; head length 9.5; head width 4.0; neck 3.0; snout-vent length 256; tail missing perhaps no more than 2���3 distal subcaudals, length 85 + N; eye diameter 1.1; naris to anterior edge of eye 3.0; head scales arranged in typical colubrid fashion with paired internasals, paired prefrontals, broad median frontal, narrow paired supraoculars, and elongate paired parietals; internasal suture 0.6, about 33 % length of prefrontal suture; frontal length 3.0, about 120 % as long as its distance from tip of snout, 67 % as long as parietals; interocular distance 3.5, 63 % occupied by frontal; mental broad, 1.5 x 1.0; two pairs of genials, subequal in size, posterior pair wing-shaped, deeply incised along median suture. Dorsal scales smooth, in 17 / 17 / 17 rows; four rows at midlength of tail; apical pits appear to be absent; supralabials 7 / 7, third and fourth entering orbit; infralabials 7 / 8, first pair in contact behind mental along midventer, first to third/ first to fourth in contact with anterior genials, respectively; third and fourth/fourth and fifth infralabials in contact with posterior genials, respectively; preoculars 1 / 1, subpreocular absent; postoculars 1 / 1; temporals 1 + 2; preventrals 2; ventrals 172; cloacal plate divided; subcaudals 63 +N (incomplete), divided; pupil round. Left maxilla bearing 12 teeth, increasing in size posteriorly to ninth tooth, with 4���5 teeth in middle of series noticeably enlarged, several posteriormost teeth smaller; tooth nine even with anterior edge of ectopterygoid process; teeth arranged in straight line except for slight medial curvature of anterior of maxilla; no tooth noticeably offset; no pronounced diastema within series. In life, black head cap extending from internasals to near posterior edges of parietals, including prefrontals, frontal, supraoculars; rostral with inverted, black U-shaped marking bordering lingual groove, bordered above with dusky white on edges of first supralabials, anterior nasals, and internasals; very small amount of beige mottling on posterior nasal, loreal and anterolateral portion of prefrontal; parietals with pale spot lying to either side of common parietal suture; lateral and posterior edges of parietals with dusky white borders; side of head including nasals, loreal, preocular, and postocular blackish; upper edges of most supralabials blackish, whitish below, with lower white area of supralabials sometimes with dark brown to blackish spotting; most of primary, secondary, and upper tertiary temporals with distinctive dark brown borders and whitish centers; mental with dark brown lateral borders; infralabials 1���5 with irregular dark brown vertical streaks either centrally or along suture; anterior one-third to onehalf of anterior genial with dark brown pigment; nuchal white collar extending from posterior edge of parietals posteriorly 2.0��� 2.5 scale lengths, about 3���4 scale lengths wide on side of head, involving posterior portion of ultimate supralabials; dorsal pigment dark brown, barely encroaching on lateral portion of ventrals, more so on subcaudals; first two rows of dorsals with whitish central dashes; dorsal scale rows 4���5 with inconspicuous brown stripe, bordered below by a darker brown line on adjacent portions of scale rows 3���4; scale row 6 with inconspicuous dark brown stripe extending to adjacent portions of scale rows 5 and 7; subcaudals white with irregular brown pigment along midventral area; distal one-third of tail mostly brown; iris dark bronze. In preservative (ethanol after formalin) dark pigment of head and body mostly dark brown; striping of body obscured; collar and ventrum white. Etymology. The specific epithet is derived from the Greek �����������ε����ς (dysmikos), meaning western, in reference to the westernmost distribution of this species of the genus. We propose standard English and Spanish names of Guerrero Slender Leaf Litter Snake and Hojarasquera Delgada de Guerrero, respectively. Distribution and habitat. The type-locality lies at 432 m asl at the edge of a small valley occupied the small village of Cuajilotla (Fig. 4) in the foothills of the eastern portion of Sierra Madre del Sur in Guerrero in tropical deciduous forest. The predominant types of vegetation at higher elevations in the region are oak forest and pine-oak forest, grading into tropical deciduous forest, which is restricted to the lowlands of foothills, valleys, and slopes (Leopold 1950). The type-locality (Fig. 5) is drained by the R��o Chapolapa, an upper tributary of the R��o Omitl��n- Azul, which flows into the R��o Papagayo. Other species of amphibian and reptiles recorded between 340���432 m in the vicinity of the type-locality were Rhinella marina, Craugastor rugulosus, Agalychnis dacnicolor, Smilisca baudinii, Anolis dunni, Aspidoscelis guttata, Holcosus undulatus, Phyllodactylus lanei, Phyllodactylus tuberculosus, Ctenosaura pectinata, Sceloporus siniferus, Urosaurus bicarinatus, Boa imperator (sensu Hynkov�� et al., 2009), and Leptodeira annulata. The taxonomic histories of snakes placed in the genus Rhadinella were most recently discussed by Myers (2011) and Campbell (2015). Most of the species in the genus occur in Nuclear Central America, with a number of them inhabiting Chiapas, Mexico. Of the species having a dark, blackish dorsum, with or without inconspicuous stripes, only two species where known from Mexico until recently: R. schistosa (Smith 1941; the generotype) from the eastern versant of Veracruz and R. posadasi from extreme southern Chiapas. The genus was seemingly absent from the Pacific versant of southern Mexico until the recent description of R. donaji Campbell (2015) from the Pacific highlands of Oaxaca. The subsequent discovery of R. dysmica at a relatively low elevation (432 m) in Guerrero is somewhat of a surprise. Snakes occurring at low elevations in tropical deciduous forest usually have rather extensive distributions in this widespread and continuous lowland habitat, not being affected by the isolating effects of elevation. Many species of Rhadinella have distinctive longitudinal body striping that serves as a useful identifier (see Myers 1974: fig. 28). The pattern in R. dysmica with the combination of dark striping on adjacent portions of scale rows 3���4 and on the sixth scale row and adjacent portions of fifth and seventh scale rows is not found in any other species of Rhadinella. Unfortunately, the dorsal patterning in the dark-colored species is often difficult to discern or obscured altogether. We are able to address the striping in R. dysmica only because of the excellent photographs taken in life. Under certain light conditions, striping pattern in life is obscured owing to the highly iridescent scales of this snake, and in preservative there is a tendency for striping to become even more obscure. The geographically closest congeners to R. dysmica are R. schistosa and R. donaji (Fig. 5). Rhadinella schistosa occurs about 300 km to the northeast, separated by the xeric Tehuac��n-Cuicatl��n Valley. Most individuals have been taken from near 1000 m asl in mesic tropical evergreen forest or cloud forest. Rhadinella donaji occurs about 275 km to the East and, similarly to R. dysmica, inhabits the Pacific versant, but their habitats are dramatically different: R. dysmica occurs in tropical deciduous forest at 432 m asl, whereas R. donaji is only known from pine-oak forest at 2195 m asl. The elevational distribution of these species remains to be ascertained but it is unlikely that their ranges overlap given habitat and elevational differences between the two species, and the fact that distribution of other species of Rhadinella are geographically restricted. There are several traits apparent in members of the genus Rhadinella with a dark dorsum. First, each species seems to be restricted to a particular habitat, whether it be cloud forest, pine-oak, rainforest, or in the case of R. dysmica, tropical deciduous forest. Secondly, the pale head pattern shows little variation within species (Myers 1974). Both of these traits provide evidence for the distinction of R. dysmica from R. donaji. The segmental counts for R. donaji and R. dysmica are high compared to other members of the genus and given their distributions it is likely they are closely related. With a sample of one for each of these species, it is not possible to address variation. Nonetheless, these species sufficiently differ from each other that their heterospecificity can hardly be doubted. Beside features outlined in the diagnosis, careful examination of these species reveals the following differences in color pattern. The most distinctive element of pattern is the bright white collar which in R. dysmica begins behind the parietals, although it is contiguous with parts of the parietals that are edged with dingy white. In R. donaji the white collar is continuous with the mostly pale parietals. In R. dysmica the temporals have mostly dull white centers with irregularly black edging and the loreal region is dark with small tan flecks, whereas in R. donaji the temporals are mostly white with irregular dark mottling and the loreal region is mostly white. The dark dorsal ground coloration barely reaches the lateral portions of the ventrals but strongly encroaches on lateral portions of the subcaudals in R. dysmica, whereas in R. donaji the dorsal coloration fails to reach the ventrals and barely encroaches on the subcaudals. In R. dysmica dark pigment along the midventer of the tail forms a grayish band, but in R. donaji the subcaudals are immaculate. The lower two scale rows in R. dysmica have very faint and irregular indications of pale dashes in their center and in R. donaji the lower fourth scale rows have distinct pale dashes which are especially conspicuous on the lower two rows., Published as part of Campillo, Gustavo, D��vila-Galav��z, Luis Fernando, Flores-Villela, Oscar & Campbell, Jonathan A., 2016, A new species of Rhadinella (Serpentes: Colubridae) from the Sierra Madre del Sur of Guerrero, Mexico, pp. 165-173 in Zootaxa 4103 (2) on pages 166-172, DOI: 10.11646/zootaxa.4103.2.6, http://zenodo.org/record/262518, {"references":["Leopold, A. S. (1950) Vegetation zones of Mexico. Ecology, 31, 507 - 518. http: // dx. doi. org / 10.2307 / 1931569","Hynkova, I, Starostova, Z. & Frynta, D. (2009) Mitochondrial DNA variation reveals recent evolutionary history of main Boa constrictor clades. Zoological Science, 26, 623 - 631.","Myers, C. W. (2011) A new genus and new tribe for Enicognathus melanauchen Jan 1863, a neglected South American snake (Colubridae: Xenodontidae), with taxonomic notes on some Dipsadinae. American Museum Novitates, 3715, 1 - 33. http: // dx. doi. org / 10.1206 / 3715.2","Campbell, J. A. (2015) A new species of Rhadinella (Serpentes: Colubridae) from the Pacific versant of Oaxaca, Mexico. Zootaxa, 3918 (3), 397 - 405. http: // dx. doi. org / 10.11646 / zootaxa. 3918.3.3","Smith, H. M. (1941) A new genus of Mexican snakes related to Rhadinaea. Copeia, 1941, 7 - 10. http: // dx. doi. org / 10.2307 / 1437694","Myers, C. W. (1974) The systematics of Rhadinaea (Colubridae), a genus of New World snakes. Bulletin of the American Museum of Natural History, 133, 1 - 262."]}

Amblystoma velasci Dugès, 1888 c There are three syntypes in the MADUG (HE 579–581, Figs 3 D, 4 A–B); two are metamorphosed and one (HE 581) is a larva. Smith & Necker (1943) did not find any specimens of this salamander; therefore they referred to the original description in “La Naturaleza” by Velasco (1879: 216), who called it Siredon Tigrina. Since that name had already been used for another species (Smith & Necker 1943), Dugès (1888 c) gave it a new name honoring J. M. Velasco, who had described it in detail based on several specimens from Lago de Santa Isabel, near Villa de Guadalupe Hidalgo, Valley of Mexico (Ciudad de Mexico). The type locality no longer exists as it has been completely urbanized by the growth of Mexico City (Ezcurra et al. 2006). The specimens in the MADUG were apparently sent by Manuel M. Villada, who was the sole editor of “La Naturaleza” from 1870 to 1914 (Cuevas Cardona 2002) (notes taken from Notebook No. 2 in the MADUG, fide GEMC). The type locality referred to by Dugès (1888 c) in his description is given as: “Laguna de Santa Isabel, en la Villa de Guadalupe y en los pueblos de Santa Isabel y Zacatengo” (the last town now called Zacatenco, Ciudad de Mexico). Current valid name: Ambystoma velasci (Dugès, 1888)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 37, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Hemichirotes tridactylus Dugès, 1894 a The original description of this species appeared in Cope (1894) in the May issue of “The American Naturalist”. Cope credited the description to Dugès’ manuscript describing a new amphisbaenian from Guerrero, Mexico. Cope published an abbreviated version of the description that Dugès published in “La Naturaleza” (Dugès 1894 b). Copes’paper reproduces five of the eight figures as they were published by Dugès (Pl. XX 1 ’ and 1–7). To satisfy the criterion of availability, according to Article 5.1. 1 of the ICZN (1999), the name should be credited to Dugès, since Cope had no specimen, at ANSP or USNM, upon which to base his description (Cochran 1961; Malnate 1971), the figures reproduced in Cope (1894) are the same as those made by Dugès except for a drawing of a scale (Figure 6 g in Cope 1894) and published later the same year. When Smith & Taylor (1950), published the checklist of Mexican reptiles; they reported that the only specimen known at the time, was the type at the MADUG (Smith & Taylor 1950: 38). The holotype for this species is still at the MADUG (HE 127; Fig. 5 B). A detailed description of this specimen is given by Smith & Necker (1943). Current valid name: Bipes tridactylus (Dugès in Cope, 1894)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on pages 39-40, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Eumeces rovirosae Dugès, 1893 The specimen at the MADUG is not in good condition, but is as Dugès reported in his original description: the specimen was “muy maltratado del vientre, cuello y ano” (very damaged on the belly, neck and anus). The holotype was described in detail by Smith & Necker (1943), and bears the number HE 189 (Fig. 6 C). Current valid name: Plestiodon sumichrasti (Cope, 1867)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 40, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Tapaya orbicularis longicaudatus Dugès, 1888 b Dugès’ original description (Dugès 1888 b) apparently was based on a single specimen. Smith & Necker (1943) reported that they found the type and gave a detailed description and measurements. In the series of Phrynosoma orbicularis specimens in the MADUG, one bears the locality “ Mexico ” and is labeled as Tapaya orbicularis and it broadly coincides with Smith & Necker’s (1943) description and measurements (HE 422, Fig. 6 A). This specimen should be considered the holotype. Current valid name: Phrynosoma o. orbiculare (Linnaeus, 1758)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 40, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Rhinocheilus antonii Dugès, 1886 This specimen is still at the MADUG, and was fully described and photographed in Smith & Necker (1943). Regarding the type locality, Dugès (1886) originally stated “…vient des environs de Mazatlán, côte du Pacifique.” (…it comes from the surroundings of Mazatlán, Pacific coast.); although Smith & Necker (1943: 210) noted that Dugès in a later description (Dugès 1888 a) mentioned “San-Blas, Mexique ”. We concur with Smith & Necker (1943) that the type locality should be San Blas, Nayarit, Mexico (see Discussion in Smith & Necker 1943: 210). The type (HE 1223, Fig. 8 B) does not have the original tag mentioned by them; it now bears a field tag from Walter L. Necker 1401. The specimen was completely dry and almost unrecognizable, recently it has been rehydrated. Current valid name: Rhinocheilus lecontei antonii Dugès, 1886., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on pages 43-44, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E., Quezadas-Tapia, Néstor L. (2016): Alfredo Dugès' type specimens of amphibians and reptiles revisited. Zootaxa 4092 (1), DOI: http://doi.org/10.11646/zootaxa.4092.1.2

Campillo, Gustavo, Dávila-Galavíz, Luis Fernando, Flores-Villela, Oscar, Campbell, Jonathan A. (2016): A new species of Rhadinella (Serpentes: Colubridae) from the Sierra Madre del Sur of Guerrero, Mexico. Zootaxa 4103 (2): 165-173, DOI: http://doi.org/10.11646/zootaxa.4103.2.6

Storeria dekayi anomala Dugès, 1888 e The type from “Orizaba (Mexique)” (Dugès, 1888 e) was redescribed and photographed by Smith & Necker (1943: 210, plate 6, fig. 10). They also reassigned the type locality to Orizaba, Veracruz. Unfortunately, the type has apparently been lost; we were unable to find it after checking all specimens belonging to the genus Storeria at the MADUG. Current valid name: Storeria dekayi anomala Dugès, 1888., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 48, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Sceloporus intermedius Dugès, 1877 In his original description, Dugès wrote that he used several specimens, at least one male that is described, a female, and an uncertain number of juvenile females based on his mention of “En las jóvenes los tintes generales…” (In juvenile females the general colors…). As usual, Dugès described a male in detail, and gave measurements for only one specimen. It is worth noting that Dugès’ SVL measurements are inconsistent since he reported “0,m04” (= 40 mm). Smith & Necker (1943) did not notice this inconsistency, but gave the measurements for two cotypes (according to them), a male and a female. Smith & Necker’s measurements do not match those given by Dugès (1877). They also stated that both specimens have incomplete tails. There are two specimens at MADUG (HE 369–370, Fig. 10 B–C), both males. The measurements for these are not the same as those given by Dugès (1877) or those given by Smith & Necker (1943) (HE 369; head 16.8 mm, body 75.9 mm, tail 99.7 mm, forelimb 28.0 mm, hind leg 41.9 mm, foot 18.9 mm). Also the specimens now at the MADUG have complete tails; that of HE 370 is broken, but complete. One of the MADUG specimens was collected at La Noria, Michoacán, the type locality given by Dugès (1877). So, specimen HE 369 can only be considered a topotype. Our search leads us to the conclusion that the specimens that Smith & Necker (1943) described are lost from the herpetological collection at MADUG. Current valid name: Sceloporus dugesii Bocourt, 1873., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 46, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Geophis tecpanecus Dugès, 1896 b The only specimen that Dugès used for this description was partially destroyed, i.e., the body in four pieces (Smith & Necker 1943). According to Dugès (1896 b) the head was smashed and almost all of the teeth and the mandibular articulation were destroyed. Dugès (1896 b) illustrated a reconstructed specimen and the portion of the skull that remained undamaged. The type specimen was still at the MADUG when Smith & Necker (1943) were there, and they presented a close up of the head. Soft parts of the type are apparently lost, and we were unable to find any indication of their existence despite checking all possible candidates. There is a skull, seven dorsal vertebra and three ribs in the collection (HE 1139, Fig. 7 B), and we assume these remains are part of the type. Dugès (1896 b) drew the skull and the specimen at the MADUG coincides with plate XXVIII and the proportions depicted on this plate. Therefore, we consider these bones to be all that remains of the holotype. There is additional information on this taxon. In 1898, Dugès described the genus Geatractus based on the same specimen arguing that the specimen presented characteristics of the genera Geophis and Atractus. He based part of his argument on the similarity of the remaining part of the skull saying it looked more similar to that of Atractus. However, he noted that the vertebrae were more similar to those of a Geophis (Dugès 1898). Current valid name: Tropidodipsas annulifera (Boulenger, 1894)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 41, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Elaps diastema michoacanensis Dugès, 1891 b As stated by Smith & Necker (1943), the original specimen that was described and illustrated in color (Dugès 1891 b: 487 – 88 + plate 32) has not been found. However, the neotype (HE 1) they designated still is at the MADUG (Fig. 8 C). A detailed description of this specimen is given in Smith & Necker (1943), and the specimen remains in good condition. It is worth pointing out that the neotype was the same specimen Dugès used in latter publications (1896 a, 1901) to elevate this taxon to the species category. Current valid name: Micrurus distans michoacanensis (Dugès, 1891)., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on page 44, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210

Phrynosoma taurus Bocourt 1870 This taxon was originally mentioned by Dugès (1869), in a list of vertebrates observed in the Republic of Mexico. Then it was illustrated and described in the “Mission Scientifique” (Bocourt, (1870), Pl. 12, figs. 8 a– 8g; (1874), pp. 234-235) with credit for the name given to Dugès (1869) and a manuscript by Dugès dated 1868. A redescription of the species was published by Dugès (1873) in “La Naturaleza”, although the published version bears the date November 23, 1870 as correctly stated by Smith & Taylor (1950). See also Smith (1942) for publication dates of “La Naturaleza”. Bocourt (1874: 234–235), published a description of this taxon, with a footnote (page 234) mentioning a manuscript written by Dugès in 1868; Dugès (1869) listing the name; and making reference, in an endnote, to a monograph of the group being published by Aug. Duméril. Bocourt’s (1874) description mentioned nine specimens (syntypes), two adults and the rest juveniles, obtained by Boucard from “…sur le plateau de Puebla au bien à Matamoros Izucar (Mexique).” (on the plateau of Puebla or in Matamoros Izucar (Mexico), at the Museum National d’Histoire Naturelle, Paris (MNHN 1270, 1915, 1915 A, 1915 B, 1916, 1310, 1310 A, 1310 B, 1310 C, fide Brygoo, 1989). Bocourt (1874) also noted that Dr. Dugès had another adult specimen from Coscatlan (Coxcatlán, Puebla, Mexico). It seems pertinent to report the accidental history of the authorship and publication date for this species. After the publications mentioning, illustrating and describing this taxon; the first review of the genus Phrynosoma was published by Gentry (1885); this author cited in the following order the publications bearing the name of this species: “ Phrynosoma taurus Dugès MSS., 1868; Dugès, Cat. Vert. Mex., 1869; Dumeril (sic) & Bocourt, Miss. Sci. au Mex. et Am. Cent. Rept., 1870, p. 234, pl. xii, fig. 8; Dugès, La Naturaleza, ii, 1871 – 73, p. 302, fig.” Reeve (1952) cited the following works: “ Phrynosoma taurus, Dugès, Desc. Manuscript (fide Duméril and Bocourt loc. cit., p. 234), 1869 (sic); and La Naturaleza, 1869, p. 143 (nomen nudum); Aug. Duméril and Bocourt, Mission Scientifique au Mexique et dans l’Amerique Centrale, Études sur les Reptiles livr. 1, 1870, pl. 12, figs. 8, 8 a–g; Bocourt, idem, livr. 4, 1874, pp. 234–235, pls. 1, 2, (sic) figs. 8 a–g; Dugès, La Naturaleza, 1873, pp. 302– 305, figs. 1–4; Boulenger, Catalogue of Lizards in the British Museum of Natural History, vol. 2, 1885, p. 249;…” Smith & Taylor (1950) cited the following: “ Phrynosoma taurus Dugès, MS. 1868, fide Duméril and Bocourt loc. cit., p. 234; and La Naturaleza, 1869, p. 143 (nomen nudum).—Aug. Duméril and Bocourt Mission scientifique au Mexique …, Études sur les reptiles, livr. 1870, pl. 12, figs. 8, 8 a–g.—Dugès, La Naturaleza, 1873, pp. 302–305, figs. 1–4 (this description bears the date Nov. 23, 1870, but was not published until 1873; there were two specimens, one from " Cozcatlán," the other from Estado de Puebla).—Bocourt, Mission scientifique au Mexique … Études sur les reptiles, livr. 4, 1874, pp. 234–235.—Boulenger, Catalogue of lizards in the British Museum, vol. 2, 1885, p. 249.—Smith and Necker Anal. Esc. Nac. Cienc. Biol., vol. 3, 1943, pp. 208–210, pl. 3, figs. 1–2.” Other sources and checklists: Guibé (1954): Phrynosoma taurus Dugès, 1874; Brygoo (1989): Phrynosoma taurus Bocourt, 1870; Flores-Villela (1993): Phrynosoma taurus Dugès, 1868 (sic); Liner & Casas-Andreu (2008): Phrynosoma taurus Dugès 1874 (1873) (sic); Uetz & Hošek, (2015) (the Reptile Database): Phrynosoma taurus Dugès, 1873; Flores-Villela et al. (2010): Phrynosoma taurus Dugès 1870. Owing to the inconsistencies related to the authorship and publication dates of this taxon, it is pertinent to point out that among the list of publications cited above, the only correct one was Brygoo (1989). He correctly credited Bocourt for the authorship of the name Phrynosoma taurus, since Bocourt had to be responsible for the publication of the text and plates, due to the premature death of Auguste Duméril on November 12, 1870, who only wrote the introduction of the book (pages 1–7). Based on Article 12.2. 7 of the ICZN (1999), the publication of plate 12 Fig. 8 a –g, in 1870 by Bocourt in Duméril and Bocourt makes the name available and credited to Bocourt, despite the fact that in the explanation of the plate, the name was credited to Dugès. The use by Bocourt of one of the nine specimens that were already housed at the Muséum National d’Histoire Naturelle, Paris, not provided by Dugès, to be illustrated on plate 12, makes Bocourt the person to satisfy the criterion of availability, according to Article 5.1. 1 of the ICZN (1999). Therefore Phrynosoma taurus Bocourt, 1870, is the correct form of citation of this species, the corrected type locality should be: Izúcar de Matamoros, Puebla Mexico. The specimen cited by Smith & Necker (1943) as a lectotype for this taxon (HE 446, Fig. 11 C) was erroneously given this status. Current valid name: Phrynosoma taurus Bocourt 1870., Published as part of Flores-Villela, Oscar, Ríos-Muñoz, César A., Magaña-Cota, Gloria E. & Quezadas-Tapia, Néstor L., 2016, Alfredo Dugès' type specimens of amphibians and reptiles revisited in Zootaxa 4092 (1) on pages 48-50, DOI: 10.11646/zootaxa.4092.1.2, http://zenodo.org/record/269210