Cryptic diversity across the Trans-Mexican Volcanic Belt of Mexico in the montane bunchgrass lizard Sceloporus subniger (Squamata: Phrynosomatidae) Author Bryson, Robert W. Department of Biology and Burke Museum, University of Washington, Box 351800, Seattle, WA 98195 - 1800, USA & Moore Laboratory of Zoology, Occidental College, Los Angeles, CA 90041, USA Author Grummer, Jared A. Department of Biology and Burke Museum, University of Washington, Box 351800, Seattle, WA 98195 - 1800, USA & Department of Zoology and Biodiversity Research Centre, University of British Columbia, Vancouver, BC, V 6 T 1 Z 4, Canada Author Connors, Elizabeth M. Moore Laboratory of Zoology, Occidental College, Los Angeles, CA 90041, USA Author Tirpak, Joseph Department of Biology and Burke Museum, University of Washington, Box 351800, Seattle, WA 98195 - 1800, USA Author Mccormack, John E. Moore Laboratory of Zoology, Occidental College, Los Angeles, CA 90041, USA Author Klicka, John Department of Biology and Burke Museum, University of Washington, Box 351800, Seattle, WA 98195 - 1800, USA text Zootaxa 2021 2021-04-19 4963 2 335 353 journal article 7135 10.11646/zootaxa.4963.2.5 795e70be-a6cd-449e-b356-d54f061dd506 1175-5326 4700962 B595034E-9246-497D-9662-21721FE595A6 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) . FIGURE 4. (A) Dorsal, (B) ventral, and (C) lateral views of the holotype of Sceloporus dixoni sp. nov. (UTA 61714) (D) Closeup view of head. FIGURE 5. Dorsal and ventral aspects of male Sceloporus dixoni sp. nov. from the type locality. Specimen photographed and released. 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. TABLE 3. Variation in morphological characters examined in male specimens of the montane bunchgrass lizard Sceloporus subniger for this study. Range with means in parentheses; italicized characters show range with mode.

Character	S. subniger (n=7)	S. subniger “West” (n=6)	S. subniger Sierra de Mascota (n=3)
Snout–Vent Length	41–60 (49.9)	42–54 (48.2)	43–47 (45)
Head Length	9.2–12.5 (10.9)	9.5–11.5 (10.3)	9.3–10.1 (9.7)
Head Width	8.3–10.8 (9.8)	8.4–10 (9.6)	8.8–9.2 (9)
Head Width / Head Length Ratio	0.85–0.98 (0.9)	0.86–0.98 (0.93)	0.91–0.94 (0.93)
Tibia Length	7–11 (8.6)	8–9 (8.8)	8–9 (8.3)
Tibia Length / Head Length Ratio	0.7–0.88 (0.79)	0.78–0.9 (0.86)	0.83–0.89 (0.86)
Dorsal Scales	39–44 (41)	40–45 (42)	42–47 (45)
Midbody Scales	34–44 (41)	39–43 (40)	40–44 (42)
Ventral Scales	27–42 (34)	32–37 (35)	38–39 (38)
Canthals	1–1 (1)	1–1 (1)	1–1 (1)
Loreals	1–2 (1)	1–1 (1)	1–2 (1)
Supralabials	4–5 (4)	4–5 (5)	4–5 (4)
Infralabials	5–6 (6)	6–7 (6)	5–7 (6)
Frontoparietals	1–2 (1)	1–2 (1)	1–1 (1)
Parietals	1–1 (1)	1–1 (1)	1–1 (1)
Scales Bordering Interparietal	14–17 (16)	13–18 (15)	11–17 (14)
2 nd Pair of Postmentals	1–2 (2)	0–2 (2)	1–2 (2)
3 rd Pair of Postmentals	3–5 (5)	2–5 (5)	4–4 (4)
Femoral Pores	32–40 (35)	31–36 (34)	34–36 (35)
Fourth Toe Lamellae	15–21 (17)	16–22 (19)	16–20 (17)

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. TABLE 4. Variation in morphological characters examined in female specimens of the montane bunchgrass lizard Sceloporus subniger for this study. Range with means in parentheses; italicized characters show range with mode.

Character	S. subniger (n=21)	S. subniger “West” (n=8)	S. subniger Sierra de Mascota (n=2)
Snout–Vent Length	41–62 (48.1)	42–52 (46.3)	45–47 (46)
Head Length	8.7–11.8 (9.8)	8.7–10.2 (9.5)	9.2–9.4 (9.3)
Head Width	7.8–10.7 (8.9)	8–9.7 (8.7)	8.4–8.8 (8.6)
Head Width / Head Length Ratio	0.85–1.04 (0.91)	0.84–0.98 (0.92)	0.9–0.96 (0.93)
Tibia Length	7–10 (8.3)	7–9 (8.3)	7–8 (7.5)
Tibia Length / Head Length Ratio	0.76–0.92 (0.85)	0.76–0.94 (0.87)	0.75–0.87 (0.81)
Dorsal Scales	37–46 (40)	37–42 (40)	41–42 (41)
Midbody Scales	37–44 (41)	37–42 (39)	43–45 (44)
Ventral Scales	31–41 (34)	32–39 (35)	37–39 (38)
Canthals	1–1 (1)	1–1 (1)	1–1 (1)
Loreals	1–2 (1)	1–2 (1)	1–1 (1)
Supralabials	3–5 (4)	4–5 (4)	4–5 (4)
Infralabials	5–6 (6)	4–6 (6)	5–6 (6)
Frontoparietals	1–2 (1)	1–2 (1)	1–1 (1)
Parietals	1–2 (1)	1–1 (1)	1–1 (1)
Scales Bordering Interparietal	14–17 (15)	12–17 (14)	15–15 (15)
2 nd Pair of Postmentals	0–3 (2)	1–2 (2)	1–2 (1, 2)
3 rd Pair of Postmentals	3–6 (5)	4–5 (4)	3–4 (3, 4)
Fourth Toe Lamellae	15–22 (18)	16–21 (19)	18–18 (18)

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 ) .