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Sceloporus dixoni sp. nov. Bryson & Grummer Figs. 4–5, Tables 3–4 Sceloporus aeneus – Duellman 1965 (in part) Sceloporus aeneus – Thomas & Dixon 1976 (in part) Sceloporus aeneus – Benabib et al. 1997 (in part) Sceloporus aeneus aeneus – Smith 1937 (in part) Sceloporus aeneus aeneus – Smith 1939 (in part) Sceloporus aeneus aeneus – Schmidt & Shannon 1947 Sceloporus aeneus aeneus – Duellman 1961 (in part) Sceloporus aeneus aeneus – Mink et al. 1996 (in part) Sceloporus aeneus subniger – Smith et al. 1993 (in part) Sceloporus aeneus subniger – Bryson et al. 2012 (in part) Sceloporus subniger – Grummer et al. 2014 (in part) Holotype: Adult male, UTA 61714 (field number RWB 0649), from Nevado de Colima, 13.5 mi W Cd. Guzmán, municipality of San Gabriel, Jalisco (N 19.6427°, W 103.6236°, 2375 m; WGS84); collected 24 June 2006 by R. W. Bryson Jr. Paratypes: same data as holotype (MZFC 22053, 22054; UTA 61713, 61715–61716). Michoacán: 11.7 mi W Zacapu on rd to Zamora (MZFC 22055, 22056; UTA 61699 –61702). 22 km N Uruapan on Hwy 37 (UTA 61703, 61704). Diagnosis. Sceloporus dixoni is a member of the S. scalaris group, sharing with other species in this group parallel lateral scale rows, femoral pore series that are either in contact or separated by no more than two scales, females with smooth preanal scales, and males with lateral abdominal color patches (Smith 1939; Smith et al. 1997; Watkins-Cowell et al. 2006). Sceloporus dixoni can be distinguished from other species in this group by the following combination of characters: single canthal on each side of the head, small adult size (maximum SVL = 54 mm, average 47.1 mm), 37–45 dorsal scales (average 41), 37–43 scales around midbody (average 40), 32–39 ventral scales (average 35), tibia length/head length proportion of 0.76–0.94 (average 0.86), 4–5 supralabial scales (mode of 5), 12–18 scales bordering the interpariatel scale (average 15), 31–36 femoral pores in males (average 34), adult females with lightly mottled venters, and adult males with extensive dark pigment on the venter, heavily mottled throats, and orange or rust-colored flanks. Description of holotype. Adult male (Fig. 4). SVL = 53 mm, total length including tail = 124 mm. Head length = 10.16 mm. Tibia length = 9 mm. Entire hind limb length (including fourth toe) = 21 mm. Forelimb length = 10.6 mm. Dorsal head scales keeled with smooth margins. Four internasal scales about twice as high as wide. Canthals 1-1. Loreals 1-1. Supralabials 5-5. Infralabials 6-6. Postnasals 3-2. Preoculars 1-1, with strong transverse keel on dorsal portion. Three frontonasals, each with>3 ridges. Three prefrontals, two large lateral ones (each with three ridges) and one small medial one with a single ridge. Frontal trapezoidal, with a medial depression and ridges on lateral portions. Frontoparietals 1-1. Parietals 2-1. Lorilabial rows 2-2. Dorsal scales triangular, keeled; about 75% of them possessing a spiny distal projection. Dorsal scale margins smooth (not serrate), transparent. Forty dorsal scales. Forty scales around midbody. Ventral scales rounded with a notch at posterior apex. Color in preservative. Dorsal and lateral surface of head medium brown. Suboculars, loreals, canthals, and labiomentals white. Throat dark blue/black with about 10 light-colored scales scattered across gular region. Dorsum medium brown, patternless. Lateral areas of body light brown and turquoise. Venter dark with turquoise in posteromedial portion, slightly less melanized near intersection with hind limbs. Dorsal surface of tail medium brown, patternless, turning to light brown towards tail tip; ventral surface cream. Forelimbs same ground color as dorsum; elbows and forearms with turquoise scales. Hindlimbs same color as dorsum. Variation. Variation in meristic and mensural characters of male and female paratypes is summarized in Tables 3–4. All males have heavily mottled throats; in several, the mottling is so dense that the ventral surface of the head appears almost entirely black, as seen in the holotype. Ventral surfaces of males are similarly dark in preservative; in some, a pale-colored patch extends midventrally from about the intersection of the hindlimbs towards the front limbs. This lighter-colored section of the venter is especially evident in life, as seen in Fig. 5. Also noticeable in this image are the lateral blue patches on the venter and orange-red color of the flanks of males. In preservative, the ventral surface darkens considerably, presumably due to fixation in formalin. The dorsal surface of males ranges from weakly patterned to patternless. When patterned, the dorsal surface is marked by a pair of light-colored dorsolateral stripes, one-scale wide, that originates at the posterior margin of ear opening and extends onto the tail. A pale vertebral line, two scale-rows wide, is also present, beginning at the nape of the neck and extending posteriorly to tail. The region between the vertebral and dorsolateral stripe is marked with narrow, dark brown transverse bars on each side; in many individuals, these bars are dimly evident. Females possess lightly mottled throats, some with more mottling than others. The ventral surface of females is very lightly mottled. The dorsal surface of females ranges from strongly patterned to patternless. In strongly patterned individuals, dark transverse bars are sharply defined, often edged posteriorly by white. Comparisons. Sceloporus dixoni is most similar to S. subniger and specimens from the Sierra de Mascota in western Jalisco, sharing with them a single canthal on each side of the head, relatively short legs (average tibia length/head length proportion less than 0.9), small adult size (maximum SVL less than 63 mm), 36–50 dorsal scales, extensive dark pigment on the venter of adult males, a black-barred or darkly mottled chin/throat in adult males, orange or rust-colored flanks in adult males, and oviparity. Sceloporus dixoni can be distinguished from S. subniger by the combination of its smaller adult size (maximum SVL = 54 mm in S. dixoni vs. 62 mm in S. subniger; average SVL = 47.1 mm vs. 48.6 mm), longer legs (average tibia length/head length proportion 0.86 vs. 0.83), fewer femoral pores in males (maximum of 36 vs. 40; average number 34 vs. 35), fewer scales around midbody (average of 40 vs. 41), more supralabial scales (mode of 5 vs. 4), and fewer scales bordering the interpariatel scale (average of 15 vs. 16). Female S. dixoni also have considerably less mottling on the ventral surface than female S. subniger. Sceloporus dixoni differs from specimens from the Sierra de Mascota in western Jalisco by the combination of their larger adult size (maximum SVL = 54 mm in S. dixoni vs. 47 mm in specimens from the Sierra de Mascota; average SVL = 47.1 mm vs. 45.4 mm), slightly longer legs (average tibia length/head length proportion 0.86 vs. 0.84), fewer ventral scales (a minimum of 32 vs. 37; average = 35 vs. 38), fewer dorsal scales (37–45, average = 41 vs. 41–47, average = 43), and fewer scales around midbody (37–43, average = 40 vs. 40–45, average = 43). Etymology. The specific epithet is a patronym honoring the late James R. Dixon for his decades of research on Mexican herpetofauna, including several insightful studies of the S. scalaris group. “Doc” Dixon took an early interest in the academic growth of the first author and made a profound and lasting impact. For this and for his encouragement and support, he will be truly missed. Distribution. Sceloporus dixoni is distributed in primarily pine-oak forest along the western half of the Trans-Mexican Volcanic Belt, from near Morelia, Michoacán, to the lower slopes of the Nevado de Colima in Jalisco. East of Morelia, the series of steep low-elevation drainages leading into the Balsas Basin likely serve as a geographic barrier between S. dixoni to the west and S. subniger to the east (Fig. 1). Comments. Several species in the S. scalaris group form a distinct subgroup based on morphology (Smith et al. 1993) and genetic data (Mink & Sites 1996; Benabib et al. 1997; Bryson et al. 2012; Grummer et al. 2014; Leaché et al. 2016), including S. aeneus Wiegmann, S. bicanthalis, S. subniger, S. dixoni, and specimens from the Sierra de Mascota in western Jalisco. Sceloporus bicanthalis is the only species in this subgroup that is viviparous and that has two vs. one canthal scales on each side of the head. Confusion regarding parity in these species was clarified by Méndez-de la Cruz et al. (1998). All species inhabit montane bunchgrass meadows along the length of the Trans-Mexican Volcanic Belt of Mexico. The taxonomic placement of S. subniger has varied since its description as a subspecies of S. aeneus (Poglayen & Smith 1958). Thomas & Dixon (1976) argued that S. a. aeneus and S. s. subniger were indistinguishable. Smith et al. (1993) challenged this conclusion, claiming it was based on misidentified specimens from Nevado de Toluca and therefore an inaccurate description of the status and distribution of S. a. subniger. Sceloporus subniger and S. aeneus were subsequently considered distinct species in checklists (Liner 1994; Bell et al. 2003), a taxonomic proposal consistent with multilocus genetic data (Grummer et al. 2014). Based on molecular data (Bryson et al. 2012; Grummer et al. 2014), the distribution of S. aeneus is certainly much smaller than envisioned by Smith in his early studies (e.g., Poglayen & Smith 1958). This smaller distribution is more accurately reflected in Smith’s later maps (e.g., Smith et al. 1993). The absence of a black-barred or mottled chin/throat and smaller adult size may distinguish S. aeneus from S. subniger (Smith et al. 1993).

Sceloporus hesperus sp. nov. Bryson & Grummer Figs. 6���7, Tables 3���4 Holotype: Adult male, MZFC 35571 (field number RWB 1107), from 2.2 km (by air) SE Lago de Juanacatl��n, Sierra de Mascota, municipality of Mascota, Jalisco (N 20.6102��, W 104.7208��, 2314 m; WGS84); collected 10 April 2011 by R. W. Bryson Jr. and M. Torocco. Paratypes: Same data as holotype (MZFC 35572���35575). Diagnosis. Sceloporus hesperus is a member of the S. scalaris group, sharing with other species in that group parallel lateral scale rows, femoral pore series that are either in contact or separated by no more than two scales, females with smooth preanal scales, and males with lateral abdominal color patches (Smith 1939; Smith et al. 1997; Watkins-Cowell et al. 2006). Sceloporus hesperus can be distinguished from other species in this group by the following combination of characters: single canthal on each side of the head, small adult size (SVL less than 47 mm, average 45.4 mm), 41���47 dorsal scales (average 43), 40���45 scales around midbody (average 43), 37���39 ventral scales (average 38), tibia length/head length proportion of 0.75���0.89 (average 0.84), 11���17 scales bordering the interpariatel scale (average 14), adult females with lightly mottled or pale venters, and adult males with extensive dark pigment on the venter, heavily mottled throats, and orange or rust-colored flanks. Description of holotype. Adult male (Fig. 6). SVL = 43 mm, total length including tail = 112 mm. Head length = 9.64 mm. Tibia length = 8 mm. Entire hind limb length (including fourth toe) = 21 mm. Forelimb length = 11 mm. Dorsal head scales keeled with smooth margins. Four internasal scales, two deeply notched towards midline and two rectangular. Canthals 1-1. Loreals 1-1. Supralabials 5-4. Infralabials 6-6. Postnasals 2-3. Preoculars 1-1 with strong transverse keel on dorsal surface. Three frontonasals, each with many ridges. Three prefrontals, each with many ridges. Frontal trapezoidal, with three ridges. Frontoparietals 1-1. Parietals 2-1. Lorilabial rows 2-2. Dorsal scales keeled, triangular, with sharp posterior point. Dorsal scale margins transparent, smooth (not serrate). Forty two dorsal scales. Forty scales around midbody. Ventral scales rounded, with a notch at posterior apex. Two white dorsolateral stripes, one-scale wide, that begin at posterior margin of ear opening and extend posteriorly to tail. Pale brown vertebral stripe two scale-rows wide. Five dark brown transverse bars between vertebral and dorsolateral stripe on each side. Gular region mostly black with about 10% of scales white, randomly scattered. Proximal dorsal surface of tail with four dark brown blotches on either side of midline; few blue scales scattered on proximal half. Color in preservative. Dorsal and lateral surface of head brown; labials dark brown, bordered dorsally and ventrally by one row of white scales. Loreal scales white. Dorsum light brown, with dark brown transverse bars and light dorsolateral stripes as described above. All white regions on dorsum infused with blue (presumably from leeching). Lateral body scales light, with increasing melanism towards venter. Throat dark blue or black, with few, scattered light-colored scales. Venter dark blue and melanized, except for midventral, pale-colored patch about six scale-rows wide extending anteriorly from inguinal region, diminishing anteriorly until absent on chest. Dorsal surface of tail as above; ventral surface cream. Limbs same color as dorsum, with few scattered light blue scales. Variation. Variation in meristic and mensural characters of male and female paratypes is summarized in Tables 3���4. All males have heavily mottled, dark turquoise throats. Ventral surfaces of two males (including the holotype) are similarly dark in preservative, with a distinct pale-colored patch extending midventrally from about the intersection of the hindlimbs towards the front limbs; in the third, the belly is much less melanized. The dorsal surface of males ranges from weakly patterned to patternless. In weakly patterned individuals, such as the holotype, the dark brown transverse bars between the vertebral and dorsolateral stripes are dimly evident. Figure 7 shows the coloration of an adult male in life; particularly noticeable are the orange-colored flanks of males. Of the two paratype females, one has a lightly mottled throat and ventral surface, while the other is pale. The dorsal surface of one female is strongly patterned, and marked with sharply defined dark transverse bars; the other female is patternless. Comparisons. Sceloporus hesperus is one of the smallest species in the S. scalaris group, having a mean SVL of 45.4 mm. Sceloporus chaneyi, previously reported to be the smallest S. scalaris group species, has a mean SVL of 45.7 mm (Liner & Dixon 1992). Sceloporus hesperus is most similar to S. subniger and S. dixoni, sharing with them a single canthal on each side of the head, relatively short legs (tibia length/head length proportion less than 0.9), small adult size (maximum SVL less than 63 mm), 36���50 dorsal scales, extensive dark pigment on the venter of adult males, a black-barred or darkly mottled chin/throat in adult males, orange or rust-colored flanks in adult males, and oviparity. Sceloporus hesperus can be distinguished from S. subniger by the combination of its smaller adult size (maximum SVL = 47 mm in S. hesperus vs. 62 mm in S. subniger; average SVL = 45.4 mm vs. 48.6 mm), more dorsal scales (average of 43 vs. 41), more scales around midbody (average of 43 vs. 41), more ventral scales (average of 38 vs. 34), and fewer scales bordering the interpariatel scale (average of 14 vs. 16). Female S. hesperus also have considerably less mottling on the ventral surface than female S. subniger. Sceloporus hesperus differs from S. dixoni by the combination of their smaller adult size (maximum SVL = 47 mm in S. hesperus vs. 54 mm in S. dixoni; average SVL = 45.4 mm vs. 47.1 mm), slightly shorter legs (average tibia length/head length proportion 0.84 vs. 0.86), more ventral scales (average = 38 vs. 35), more dorsal scales (41���47, average = 43 vs. 37���45, average = 41), and more scales around midbody (40���45, average = 43 vs. 37���43, average = 40). Etymology. The specific epithet is derived from the Greek word hesperos, meaning ���western���, and is used in reference to the type locality located at the far western end of the Trans-Mexican Volcanic Belt. Distribution. Smith (1939: 356) cites several published records from the late 1800s for S. aeneus aeneus from Jalisco: ���N of Rio Santiago (G��nther 1890); La Cumbre de los Arrastrados (Boulenger 1897); Hda. Santa Gertrudis (Boulenger 1897).��� Over 50 years later, Smith et al. (1993: 133) commented that Boulenger���s records from Jalisco ���appear to be too far west to be accepted without verification.��� If Boulenger���s records are indeed correct, specimens from La Cumbre de los Arrastrados and Hacienda Santa Gertrudis in the Sierra de Cuale might be referable to S. hesperus. A narrow ridge above 2,200 m extends across southwestern Jalisco from the type locality in the Sierra de Mascota through the Sierra de Cuale to the Sierra de Manantl��n. The reptiles and amphibians in this remote region of Mexico remain poorly sampled (Reyes-Velasco et al. 2010). If suitable montane bunchgrass habitat is present along the forested mountain pass connecting these three sierras, then it is conceivable that additional populations of S. hesperus will be found here. For now, the only known population of S. hesperus is from the high-elevation pineoak forest at the type locality in the Sierra de Mascota of Jalisco. The low-elevation valleys trending northwest from the Nevado de Colima likely serve as a geographic barrier between S. hesperus and S. dixoni. This lower-elevation area is also inhabited by S. unicanthalis and S. scalaris (Thomas & Dixon 1976; Watkins-Colwell et al. 2006), two larger-bodied species in the S. scalaris group not known to co-occur with S. subniger., Published as part of Bryson, Robert W., Grummer, Jared A., Connors, Elizabeth M., Tirpak, Joseph, Mccormack, John E. & Klicka, John, 2021, Cryptic diversity across the Trans-Mexican Volcanic Belt of Mexico in the montane bunchgrass lizard Sceloporus subniger (Squamata: Phrynosomatidae), pp. 335-353 in Zootaxa 4963 (2) on pages 348-350, DOI: 10.11646/zootaxa.4963.2.5, http://zenodo.org/record/4700962, {"references":["Smith, H. M. (1939) The Mexican and Central American lizards of the genus Sceloporus. Zoological Series: Field Museum of Natural History, 26, 1 - 427.","Smith, H. M., Watkins-Colwell, G. J., Lemos-Espinal, J. A. & Chiszar, D. (1997) A new subspecies of the lizard Sceloporus scalaris (Reptilia: Sauria: Phrynosomatidae) from the Sierra Madre Occidental of Mexico. Southwestern Naturalist, 42, 290 - 301.","Liner, E. A. & Dixon, J. R. (1992) A new species of the Sceloporus scalaris group from Cerro Pena Nevada, Nuevo Leon, Mexico (Sauria: Iguanidae). The Texas Journal of Science, 44, 421 - 427.","Gunther, A. C. L. G. (1890) Biologia Centrali-Americana. Reptilia and Batrachia. Porter, London.","Boulenger, G. A. (1897) A revision of the lizards of the genus Sceloporus. Proceedings of the Zoological Society of London, 1897, 474 - 522.","Smith, H. M., Camarillo-Rangel, J. L. & Chiszar, D. (1993) The status of the members of the Sceloporus aeneus complex (Reptilia: Sauria) of Mexico. Bulletin of the Maryland Herpetological Society, 29, 130 - 139.","Reyes-Velasco, J., Grunwald, C. I., Jones, J. M. & Weatherman, G. N. (2010) Rediscovery of the rare Autlan longtailed rattlesnake, Crotalus lannomi. Herpetological Review, 41, 19 - 25.","Thomas, R. A. & Dixon, J. R. (1976) A re-evaluation of the Sceloporus scalaris group (Sauria: Iguanidae). Southwestern Naturalist, 20, 523 - 536.","Watkins-Colwell, G. J., Smith, H. M. & Chiszar, D. (2006) Sceloporus scalaris Wiegmann. Catalogue of American Amphibians and Reptiles, 814.1 - 814.10."]}