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The spiny lizard genus Sceloporus was described by Wiegmann in 1828, with S. torquatus posteriorly designated as the type species. The taxonomic history of S. torquatus is complicated, as it has been confused with other taxa by numerous authors. Many modern systematics works have been published on Sceloporus, but none have included all five recognized S. torquatus subspecies: S. t. torquatus, S. t. melanogaster, S. t. binocularis, S. t. mikeprestoni, and S. t. madrensis. Additionally, there is previous evidence for at least one unnamed taxon. The present study is the first taxonomic revision of the enigmatic S. torquatus based on molecular phylogenies using combined molecular data from 12S, ND4 and RAG1 genes, and Maximum Likelihood and Bayesian inference phylogenetic methods. This work includes the most extensive sampling across the entire distribution, as well as divergence time estimates and environmental niche modelling, which combined offer a spatio-temporal framework for understanding the evolution of the species. Additionally, a series of morphological characters are analyzed to identify significant differences between lineages consistently recovered in the molecular phylogenies. Using this integrative approach, evidence is presented for eight lineages within the S. torquatus complex, five of which correspond to previously recognized subspecies and three represent unnamed taxa masked by morphological conservatism. Finally, to maintain taxonomic stability a lectotype and paralectoype are designated for S. torquatus, and certain taxonomic changes are suggested in order to reflect the phylogenetic relationships within the S. torquatus complex.

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."]}