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Sarcohyla toyota sp. nov. Toyota’s Treefrog, Rana arborícola de Toyota (Figs. 1, 2, 3, 4A, 4D, 4G, 4J, 5A, 5D, 5G, 6, 10A, 10B) Hyla thorectes— Duellman (2001): 886 –887, 1132, in part. Plectrohyla sp. 5 aff. P. thorectes— Faivovich et al. (2005): 35, 66, 71, 103, 183. Plectrohyla aff. thorectes sp5 — Wiens et al. (2010): 881. Plectrohyla thorectes— Caviedes-Solis (2013): 4, 8, 20, 29, 32, 34, 39, in part. Plectrohyla aff. thorectes— Kaplan et al. (2016): 265. Sarcohyla aff. thorectes— Caviedes-Solis & Nieto-Montes de Oca (2018): 8,13,17. Sarcohyla sp. — Faivovich et al. (2018): 4. Holotype. MZFC 34666 (CIG-1066) (Fig. 1 in life; Fig. 2 in preservative), an adult male from Mexico: Guerrero: 11.4 km (by road) SW of Puerto de Gallo, Municipio de Atoyac de Álvarez, 2020 masl, collected 11 September 2016 by C. I. Grünwald, H. Franz-Chávez, and K. I. Morales-Flores. (Figs. 1, 2) Paratypes (5). MZFC 34662–65 (CIG- 1062–1065) (Figs 3 B–3E, 4D, 4J, 5A, 5D), topotypic, with same data as type specimen; MZFC 34661 (CIG-0924) (Figs 3A, 4A, 4G), topotypic, collected 15 July 2016 by C. I. Grünwald and H. Franz-Chávez. Diagnosis. Small, broad-headed Sarcohyla (males 28.5–32.2 mm) with distinctly short snout, long fingers, long legs and head broader than body. Adult males to 32.2 mm SVL. Adult female size unknown. Eyes distinctly large, tympanum distinctly small (TW/SVL ratio 0.03), and snout short (END/HL ratio 0.23–0.27). Numerous tubercles beneath cloacal opening (Fig. 5G). Venter white with dark gray and brown marbling (Figs. 2B, 4G, 4J) and lacking any iridescent green coloration on dorsum (Figs. 4A, 4D). Comparisons with similar species, with character states of Sarcohyla toyota in parenthesis: Sarcohyla thorectes (Figs. 4B, 4E) differing by presence of iridescent green blotches on dorsum (complete absence of iridescent green; Figs. 4A, 4D); white coloration with dark gray and brown marbling on anterior 60% of venter, including chest and abdomen, posterior 40% of venter flesh-colored (Figs. 4H, 4K) (white or yellow coloration with dark gray and brown marbling on 100% of venter, no flesh colored skin present; Figs. 4G, 4J); distinct rostral keel present (Fig. 5A) (rostral keel absent; Fig. 5B); dark brown markings on face and flank not continuous, not forming mask (Fig. 5E)(dark brown markings and face and flank continuous, forming mask; Fig. 5D); tympanum distinct,>26% of ED (tympanum indistinct, ≤23% of ED); nostrils slightly protruding, head bluntly rounded in dorsal profile (Fig. 5B) (nostrils greatly protruding, head distinctly truncate in dorsal profile; Fig. 5A); lateral profile of snout rounded (lateral profile of snout distinctly truncate); yellow ventral coloration completely lacking or limited (Figs. 4H, 4K) (varying amounts of yellow ventral coloration present on throat, chest, sides of abdomen, thighs and feet; Figs. 4G, 4J); superarticular tubercles on first phalange of fingers on hand conical or barely bifid (superarticular tubercles on first phalange of fingers hand distinctly bifid); tarsal fold distinct (tarsal fold indistinct or absent); subarticular tubercles on foot conical (subarticular tubercles on foot slightly bifid); outer metatarsal tubercle absent or indistinct (outer metatarsal tubercle distinct). Sarcohyla toyota further differs from S. thorectes by following mensural proportions (values for S. thorectes given in parenthesis): longer hand, HA/SVL ratio 0.32–0.37 (0.27–0.33); smaller tympanum, TW/SVL ratio in males 0.027 –0.029 (0.035 –0.039); larger eye ED/SVL ratio 0.13–0.14 (0.12–0.13); lower TW/ED ratio in males 0.20–0.23 (0.27–0.32); shorter snout END/HL ratio 0.23–0.27 (0.27–0.29). Sarcohyla hazelae differing by (values for S. toyota given in parenthesis): larger body size in males 35.0– 38.8 mm SVL (28.5–32.2 mm SVL); inmaculate white venter (Figs. 4I, 4L) (white or yellow background with dark gray and brown marbling on 100% of venter; Figs. 4H, 4K); green dorsal ground coloration with iridescent green spots (Figs. 4C, 4F) (no green ground coloration or iridescent green spots present on dorsum; Figs. 4A, 4D); snout longer and rounded in both dorsal and lateral profile (Figs. 5C, 5F) (short snout truncate in both dorsal and lateral profile; Figs. 5A, 5D); superarticular tubercles on first phalange offingers on hand conical or barely bifid (superarticular tubercles on first phalange of fingers hand distinctly bifid). Sarcohyla toyota further differs from S. hazelae by the following mensural proportions (values for S. hazelae given in parenthesis): longer hand, HA/SVL ratio 0.32–0.37 (0.30–0.32); smaller tympanum, TW/SVL ratio in males 0.027 –0.029 (0.035 –0.038); larger eye, ED/SVL ratio 0.13–0.14 (0.10–0.11); lower TW/ED ratio in males 0.20–0.23 (0.33–0.37); shorter snout END/HL ratio 0.23–0.27 (0.27–0.28). Description of holotype. Adult male, 29.8 mm SVL; head broad and wider than body; top of head flat; HL 10.2 mm; HW 9.7 mm; head longer than wide, cephalic index 1.05; HL/SVL ratio 0.34; nostrils protuberant, oval, directed horizontally; snout truncate in dorsal profile with no distinct rostral keel (Fig. 1B); snout short and truncate in lateral profile (Fig. 1A); canthus moderately angular; loreal region concave; lips thick and barely flared; ED 4.0 mm; ELW 2.4 mm; IOD 5.7 mm; IND 2.4 mm; END 2.7 mm; ETD 2.1 mm; tympanum small, round and indistinct, TW 0.8 mm; TW/ED ratio 0.2; supratympanic fold from eye, above tympanum and down to above arm insertion, barely covers upper edge of tympanum; HaL 9.7 mm; fingers long, 3FL 6.8 mm, 4FL 4.9 mm, 3FL/SVL ratio 0.23; finger lengths 1 I 3-3 II 3-3 III 2 ⅘-2¾ IV; finger discs large, 3FPW 1.9 mm, 3FPW/3FW ratio 2.1; 4FPW 1.8 mm, 4FPW/4FW ratio 1.9, 4FPW/TW ratio 2.3; prepollex enlarged, bifid, covered with nuptial excrescenses consisting of minute spinules on prepollex and thumb; subarticular tubercles large, bifid; indistinct supernumerary pustules rounded; palmar tubercle bifid; dermal fold on wrist; UaL 6.4 mm; FaL 7.1 mm; tubercles on ventrolateral edge of forearm in row, interrupted, not forming a continuous fold; FL 14.2 mm; toes long, 3TL 6.7 mm, 4TL 8.9 mm, 4TL/SVL ratio 0.28; toe lengths 1I 2 ½-2½ II 1 ½-2¾ III 2 ½-2½ IV 2-1 ¾ V; toe discs large, 4TPW 1.7 mm, 4TPW/4TW ratio 1.8, 4TPW/TW ratio 2.1; toe subarticular tubercles moderately large, round, not bifid: indistinct supernumerary pustules rounded; inner metatarsal tubercle distinct, moderately rounded; outer metatarsal tubercle present, small, rounded in shape; low tarsal fold present but indistinct; a row of slightly raised tubercles on tarsus; FeL 13.4 mm; TL 15.1 mm; TaL 8.3 mm; tibio-tarsal articulation extends to anterior portion of eye; cloacal opening directed postero-ventrally; cloacal sheath moderately long, grooved medially; no transverse fold above cloacal opening; six large pale yellow tubercles on posterior surface of thigh and below level of cloacal opening. Skin on dorsum smooth; lateral skin shagreen; skin on venter of head shagreen, ventral skin of body and thighs granular. No axillary membrane. Dentigerous processes of prevomers large and postero-medially inclined between two moderate-sized ovoid choanae. Vocal slits present, large, extending from median part of tongue to angle of jaw. Vocal sac single, median, subgular and moderately distensible. Pupil horizontal. Color of holotype in life: Dorsum of head, body and limbs yellowish brown, mottled with indistinct blotches of darker brown; no iridescent green spots or blotches present; flanks bright yellow; pale brown blotches present on snout and lips; continuous dark brown dorso-lateral stripe extending from behind eye through tympanum and along flank to groin, equally dark brown blotches on flank above brown dorso-lateral stripe; no inter-orbital bar; no barring on limbs (Figs. 1A, 1B, 1C). Lower portions of head and throat pale yellow; chest and abdomen white and yellow with bold dark gray and brown marbling on entire chest and abdomen, with no flesh colored skin present; ventral coloration of limbs yellowish-orange. Iris coppery brown. Color of holotype in preservative: Dorsum of head and body gray with dark brown indistinct blotches, dorsal surfaces of limbs beige; flanks gray with dark brown marbling; venter white to cream, with dark gray marbling; throat white; ventral surfaces of limbs beige (Figs. 2A, 2B). Variation. The morphometric variation of the type series is summarized in Table 2. Variation in color as follows: four specimens with yellow or yellowish brown dorsal coloration, one tan, one darker brown with yellow highlights; varying degrees of dark gray and brown marbling on venter; three paratypes with abundant yellow coloration on venter, three with sparse yellow ventral coloration (Figs. 3 A–3E); 6–13 tubercles present below cloacal opening. Advertisement call. Male advertisement call consisting of single two–three pulse note sounding like rapid “crehk-crehk-crehk”, with length about 17–20 milliseconds repeated about every 5 seconds in call group (Fig. 7). Call groups may consist of 3–6 notes, however due to relatively low volume of call and tendency to call from above fast-flowing water it is difficult to discern how many different individuals are calling at a given time. The call has a dominant frequency of 4.75 kHz. The call is only moderately loud, and can be heard over the sound of the fast flowing water. Distribution and habitat. This species appears to be restricted to the immediate vicinity of the type locality in the central portion of the Sierra Madre del Sur of Guerrero, Mexico (Fig. 8). It has been collected between 1975–2185 masl. The habitat at tht type locality is cloud forest, with tropical evergreen forest at lower elevations and pine forest at immediately higher elevations(Fig. 9A). It has been collected only in the immediate vicinity of slow to fast moving streams, secondary to larger water ways, and with heavily vegetated banks (Fig. 9B). The frogs are usually found on vegetation above or near the water. ......continued on the next page Etymology. The specific epithet is a an invariable name in aposition honoring the Japanese automotive manufacturer Toyota, whose various four wheel drive models such as the Toyota Tacoma and the Toyota 4runner have been paramount to our field team’s access into hard-to-reach areas of the Mexican back country. Referred Specimens. Plectrohyla aff. thorectes Mexico: Guerrero: El Molote, Municipio de Atoyac de Álvarez (ANMO 020246); 8.21 km SW of Puerto de Gallo (UTA-A 56442); Carretera Nueva Dehli-La Guitarra (UTA-A 57681 (JAC 022224))., Published as part of Grünwald, Christoph I., Franz-Chávez, Hector, Morales-Flores, Karen I., Ahumada-Carrillo, Ivan T. & Jones, Jason M., 2019, A rare new treefrog of the genus Sarcohyla (Anura: Hylidae) from Guerrero, Mexico, pp. 345-364 in Zootaxa 4712 (3) on pages 347-355, DOI: 10.11646/zootaxa.4712.3.2, http://zenodo.org/record/3586326, {"references":["Duellman, W. E. (2001) The hylid frogs of Middle America. Vol. 1 & 2. Society for the Study of Amphibians and Reptiles, Ithaca, NY, 603 pp. & 1159 pp., 92 pls.","Faivovich, J., Haddad, C. F. B., Garcia, P. C. A., Campbell, J. A. & Wheeler, W. C. (2005) Systematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revision. Bulletin American Museum Natural History, 294. https: // doi. org / 10.1206 / 0003 - 0090 (2005) 294 [0001: SROTFF] 2.0. CO; 2","Wiens, J. J., Kuczynski, C. A., Hua, X. & Moen, D. S. (2010) An expanded phylogeny of treefrogs (Hylidae) based on nuclear and mitochondrial sequence data. Molecular Phylogenetics and Evolution, 55, 871 - 882. https: // doi. org / 10.1016 / j. ympev. 2010.03.013","Caviedes-Solis, I. W. (2013) Filogenia molecular del grupo Plectrohyla bistincta (Anura: Hylidae). Master's thesis. Universidad Nacional Autonoma de Mexico, Mexico, D. F., 50 pp.","Kaplan, M., Heimes, P., Zarza, E. & McCormack, J. E. (2016) On the morphology of Plectrohyla chryses (Anura: Hylida; Hylini), with comments on some controversial characters, phylogenetic relationships, and diagnosis of this species. Caldasia, 38 (2), 257 - 273. https: // doi. org / 10.15446 / caldasia. v 38 n 2.60921","Caviedes-Solis, I. W. & Nieto-Montes de Oca, A. (2018) A multilocus phylogeny of the genus Sarcohyla (Anura: Hylidae), and an investigation of species boundaries using statistical species delimitation. Molecular Phylogenetics and Evolution, 118, 184 - 193. https: // doi. org / 10.1016 / j. ympev. 2017.09.010"]}

Bryson, Robert W., Linkem, Charles W., Dorcas, Michael E., Lathrop, Amy, Jones, Jason M., Alvarado-Díaz, Javier, Grünwald, Christoph I., Murphy, Robert W. (2014): Multilocus species delimitation in the Crotalus triseriatus species group (Serpentes: Viperidae: Crotalinae), with the description of two new species. Zootaxa 3826 (3): 475-496, DOI: http://dx.doi.org/10.11646/zootaxa.3826.3.3

Crotalus tlaloci sp. nov. Figs. 4–5, Table 3 Crotalus triseriatus triseriatus — Campbell & Lamar (2004): 527 (Map 90), 593 (in part). Crotalus triseriatus triseriatus — Flores-Villela & Hernández-García (1989): 16. Crotalus triseriatus triseriatus — Pérez-Ramos et al. (2000): 34. Crotalus triseriatus triseriatus — Flores-Villela & Hernández-García (2006): 270. Holotype. Adult female (MZFC 3666) collected 20 June 1986 by Efrain Hernández-García in “Los Llanos” (18 ° 36 ’N, 99 ° 37 ’W; 2200–2300 m above sea level; asl hereafter), 10 km by road from Taxco to Tetipac, Sierra de Taxco, municipality of Tetipac, state of Guerrero, Mexico. Paratypes. 11 specimens. MEXICO: GUERRERO: Cerro del Huizteco, Sierra de Taxco, municipality of Taxco (18 ° 36 'N, 99 ° 36 'W; 2300–2520 m asl); collected 22–23 August 1986 by E. Hernández-García (MZFC 3664–3665). “Arroyo las Damas”, Sierra de Taxco, municipality of Tetipac (18 ° 38 'N, 99 ° 37 'W; 1600–1850 m asl); collected by E. Hernández-García (MZFC 3666). ESTADO DE MÉXICO: Acatitlán, municipality of Valle de Bravo; collected 7 September 1988 by T. Hentschel-Maida (MZFC 4324). Los Álamos, municipality of Valle de Bravo (19 ° 11 ' 20.2 "N, 100 °03' 57.2 "; 2201 m asl; NAD 27 Mexico); collected 23 May 2008 by J. Jones, C. I. Grünwald, and R. W. Bryson Jr. (HINIRENA 725–726). Los Álamos, municipality of Valle de Bravo (19 ° 11 ' 20.2 "N, 100 °03' 57.2 "; 2201 m asl; NAD 27 Mexico); collected 22 July 2009 by R. W. Bryson Jr. and M. Torocco (MZFC 25114–25115). MORELOS: Km 12, Carr. Cuernavaca-Ocuilán, municipality of Cuernavaca; collected 17 March 1990 by M. Torres Chávez (MZFC 4657). Carr. Cuernavaca-Ocuilán, near state border, municipality of Cuernavaca (18 ° 58 ' 54.43 "N, 99 ° 18 ' 20.43 "W; 2268 m asl; WGS 84); collected 13 June 2009 by J. Jones, C. I. Grünwald, and R. W. Bryson Jr. (MZFC 25111). MICHOACÁN: N Arroyo Seco, municipality of Aporo (19 ° 40 ' 28.3 "N, 100 ° 22 ' 35.8 "W; 2463 m asl; NAD 27 Mexico); collected 24 May 2008 by J. Jones, C. I. Grünwald, and R. W. Bryson Jr. (HINIRENA 724). Diagnosis. Crotalus tlaloci can be distinguished from all members of the C. triseriatus species group by the combination of the following characters: (1) presence of intercanthals, (2) undivided upper preocular, (3) 152–164 ventrals in males, 156–165 in females, (4) 27–33 subcaudals in males, 22–32 in females, (5) small rattle (proximal rattle width 11.1–14.5 % of head length), (6) long tail (8.9–11.3 % of total body length in males, 8.0– 10.7 % in females), (7) usually two pairs of symmetrical, similarly sized intercanthals, and (8) dark postocular stripe that noticeably narrows before reaching the posterior of the eye. Crotalus tlaloci is most similar to species of the C. triseriatus group distributed along the Trans-Mexican Volcanic Belt. Crotalus tlaloci is distinguished from these species by the presence of intercanthals (lacking in C. pusillus), undivided upper preocular (divided 14.3 % of the time in C. armstrongi and 9.4 % of the time in C. triseriatus), high number of ventrals (overlapping with C. pusillus, but mean number in C. tlaloci higher than C. pusillus: 156 vs. 154, respectively), high number of subcaudals (mean number in females higher than in C. pusillus, C. armstrongi, and C. triseriatus: 28 vs. 26, 24, and 24; in males, higher than in C. armstrongi and C. triseriatus: 30 vs. 28 in both), proportionately small proximal rattle (mean width smaller than in C. armstrongi and C. triseriatus: 12.8 % of head length vs. 14.0% and 15.8 %), and proportionately longer tail (mean length in females higher than in C. pusillus, C. armstrongi, and C. triseriatus: 9.2 % of total length vs. 8.6 %, 8.3 %, and 8.3 %). Most specimens (10 / 12) possess two pairs of symmetrical, similarly sized intercanthals, creating the appearance of butterfly wings. Of the 100 specimens in the C. triseriatus group that we examined, this symmetrical paired arrangement of intercanthal scales in the prefrontal region (“butterfly wings”) was observed in only one other specimen (C. armstrongi, CNAR 4498). Crotalus tlaloci also possess a dark postocular stripe that noticeably narrows before reaching the posterior of the eye. In C. pusillus, C. armstrongi, and C. triseriatus, the postocular stripe is generally of uniform width, although on rare occasions in C. pusillus and C. armstrongi it tapers slightly before reaching the eye. Description of the holotype. Rostral broader than high (4.0 x 2.8 mm); two internasals, in medial contact, wider than long, rectangular, convex through center of scale; two canthals, large, convex, separated by two pairs of square intercanthals; four large intersupraoculars posterior to intercanthals, followed by multiple rows of small intersupraoculars. Naris centered between prenasal and postnasal scales, prenasal larger than postnasal and wrapping around anterior aspect of rostrum; two loreals, lower larger; small upper loreal between canthal, internasal, postnasal, and lower loreal. Loreal pit midway between eye and naris, below line from middle of eye to naris, bordered by single prelacunal, postlacunal, lower preocular, and lower loreal; prelacunal contacting second and third supralabials and a single prefovial (two prefovials on left); two prefovials on right, three on left; single postfovial contacting postlacunal, lower preocular, first subocular, and fourth supralabial. Two preoculars, upper large and convex, contacting supraocular and canthal, lower preocular thin and long; three suboculars, anterior largest and in contact with fourth supralabial; three interoculabials posterior to anterior subocular; two postoculars, dorsal twice as large as ventral. Supralabials 12 / 12; infralabials 13 / 13; first infralabials in medial contact posterior to triangular mental; genials together resemble a heart shape. Midbody dorsal scale rows 22–23; preventral single; ventrals 162; subcaudals 27, last subcaudal row divided into three scales; nine rattle fringe scales; tail bearing four rattle segments. Coloration significantly faded in preservative with body blotches and bands difficult to discern from ground color. Ground color gray; occasional black speckling along body. Blotches faded to ground coloration in many places, approximately 38 blotches visible. Blotches bordered by lighter coloration than ground color, also heavily faded. Ground color of head gray, heavily stippled with dark brown throughout dorsal and anterior lateral regions of the head; rostral and supralabials 1–4 heavily stippled, supralabials 5–7 with lighter stippling in the scale center; infralabials with brown stippling on scale margins with white scale centers, extending to mouth; gular scales lightly stippled, decreasing in frequency towards the midline of the head. Ventral scutes evenly stippled, increasing in intensity posterior to mid-body with scutes becoming almost completely brown by the tail. Six dark tail bands, the first two bordered by white bands; caudal scales dark brown. Proximal rattle black, distal sections brownish. Color in life. Color in life varies (Fig. 5), with most specimens vibrantly colored, although one is darkly pigmented (MZFC 25111; Fig. 5). The holotype is known only from the preserved specimen. Variation. Two specimens lack symmetrical paired intercanthals. The first, HINIRENA 725, has one pair of anterior intercanthals but only a single large posterior intercanthal. The second, HINIRENA 724, has one pair of anterior intercanthals, one of which is subdivided, followed by a single large posterior intercanthal. Supralabials in five specimens (MZFC 3664, 3665, 4657, 25111, 25114) are horizontally divided. This split was in the 7 th, 8 th, or 9 th supralabial and when present, occurred in supralabials on both sides of the head. Juveniles (MZFC 3664, 3665, 3667, HINIRENA 726) have cream-colored proximal rattles and pale-colored tails ventrally, typical of juveniles in other species in the C. triseriatus group. Variation in meristic, morphometric, and color pattern characters within the type series is listed in Table 3. Etymology. This species is named for Tláloc, the Aztec god of rain. Habitat and distribution. Crotalus tlaloci inhabits open areas in cloud forest and humid oak-pine forest along the lower slopes of the Trans-Mexican Volcanic Belt. Although one record in the Sierra de Taxco (“Arroyo las Damas”) is at 1850 m (Flores-Villela & Sánchez-H 2003), most records are at around 2000–2400 m asl. This species is known from the states of Guerrero, Estado de México, Michoacán, and Morelos, and may range into western Puebla. The vegetation where C. tlaloci is found is characterized by broad-leaf oaks, such as Quercus candicans and Q. laurina, and dense undergrowth (Fig. 6), and is distinctly different than the drier pine-oak forest inhabited by C. triseriatus. The distribution of C. tlaloci overlaps the ranges of two alligator lizards, Barisia herrerae Zaldívar-Riverón & Nieto-Montes de Oca 2002 and B. rudicollis (Wiegmann 1828) (Zaldívar-Riverón & Nieto-Montes de Oca 2002). Interestingly, both of these alligator lizards occur in similar humid forest habitat at elevations of 2000–2500 m asl, and appear ecologically isolated from B. imbricata (Wiegmann 1828), which inhabits the surrounding drier pine-oak forest (Zaldívar-Riverón & Nieto-Montes de Oca 2001, 2002). Specimens of C. tlaloci are generally found in rocky open forest breaks and edges of cloud or humid oak-pine forest. However, we found an adult gravid female and juvenile (HINIRENA 725, 726) under logs in a clearing relatively devoid of rocky habitat.

Crotalus campbelli sp. nov. Figs. 7–8, Table 3 Crotalus triseriatus — Boulenger (1896): 581 (in part). Crotalus triseriatus armstrongi — Dorcas (1992): 87 (in part). Crotalus triseriatus armstrongi — Bryson et al. (2011): 699 (in part). Crotalus triseriatus armstrongi — Reyes-Velasco et al. (2009): 118. Crotalus armstrongi — Reyes-Velasco et al. (2013): 528 (in part). Holotype. Adult female (KU 73649) collected 25 October 1962 by Percy L. Clifton (field number PLC 3216) in the Sierra de Cuale, 9 km N El Teosinte, municipality of Talpa de Allende, state of Jalisco, Mexico. Paratypes. 5 specimens. Mexico: JALISCO: same collection data as holotype (KU 73650). Las Playitas, Las Joyas, Sierra de Manantlán, municipality of Autlán de Navarro; collected September 1985 by E. Fanti-Echegoyen (UTA R- 16352). Las Joyas, Sierra de Manantlán, municipality of Autlán de Navarro; collected September 1985 by E. Fanti-Echegoyen (UTA R- 16353). ca. 25 km SE Autlán, ca. 2.1 km (by dirt road) SE Manantlán; collected 27 July 1975 by G. M. Tilger and R. G. Arndt (AMNH R- 113191). Lago de Juanacatlán, Sierra de Mascota, municipality of Mascota (20 ° 37 ' 30.94 "N, 104 ° 43 ' 36.30 "W; 2009 m asl; WGS 84); collected 10 April 2011 by R. W. Bryson Jr. and M. Torocco (MZFC 28669). Diagnosis. Crotalus campbelli can be distinguished from all members of the C. triseriatus species group except C. armstrongi by the combination of the following characters: (1) presence of intercanthals, (2) infrequently divided upper preocular (9.1 % of the time), (3) 150–154 ventrals in males, 147–152 in females, (4) 31–32 subcaudals in males, 22–26 in females, (5) small rattle (proximal rattle width 11.0– 14.6 % of head length), (6) long tail (9.1 –11.0% of total body length in males, 7.5–8.9 % in females), (7) pale interspaces between dorsal and lateral blotches, (8) heavy venter mottling, (9) dark proximal rattle and underside of tail, and (10) usually a single large anterior intercanthal. Crotalus campbelli can be distinguished from C. armstrongi based on higher mean number of ventrals (152 in males and 149 in females vs. 141 and 144), higher mean number of subcaudals in males (31 vs. 28), less frequently divided upper preocular (9.1 % vs. 14.3 %), proportionately longer tail in males (10.3 % of total body length vs. 9.7 %), smaller mean proximal rattle width (13.0% of head length vs. 14.0%), higher mean number of dorsal blotches (48 vs. 42), and higher number of tail bands (mode of 9 vs. 6). Crotalus campbelli is most similar to members of the C. triseriatus group distributed along the Trans-Mexican Volcanic Belt, including C. pusillus, C. armstrongi, C. triseriatus, and C. tlaloci. Crotalus campbelli is distinguished from C. pusillus by possessing intercanthals and an infrequently divided upper preocular, from C. tlaloci by having an infrequently divided upper preocular, variable number of intercanthals, fewer ventrals (in females, 147–152 vs. 156–165; in males, mean number 152 vs. 156), lower mean number of subcaudals in females (24 vs. 28), proportionately shorter tail in females (8.2 % of total length vs. 9.2 %), and higher mean number of dorsal blotches (48 vs. 43), and from C. triseriatus by a higher number of ventrals (in males, 150–154 vs. 134–146; in females, mean number higher: 149 vs. 142), higher mean number of subcaudals in males (31 vs. 28), proportionately smaller proximal rattle (13.0% of head length vs. 15.8 %), and by having pale interspaces between dorsal and lateral blotches. Crotalus campbelli is most similar in general appearance to C. armstrongi, but can be distinguished from this species by characters mentioned above. Half of the specimens of C. campbelli also possess a single, large anterior intercanthal. This scale arrangement is rarely seen in C. armstrongi and C. triseriatus. Crotalus campbelli is easily distinguished from C. ravus by the lack of large head plates in the parietal region. Description of the holotype. Rostral broader than high (3.7 x 2.6 mm); two internasals, in medial contact, slightly wider than long, convex through center of scale; two canthals, large, convex, circular, separated by a single, large, convex intercanthal bordered anteriorly by three small scales; four large intersupraoculars posterior to intercanthals, followed by multiple rows of small intersupraoculars. Nostril centered between prenasal and postnasal scales, prenasal larger than postnasal and wrapping around anterior aspect of rostrum; single loreal. Loreal pit midway between eye and naris, below line from middle of eye to naris, bordered by single prelacunal, postlacunal, lower preocular; prelacunal contacting second and third supralabials and two prefovials; three prefovials; single postfovial contacting postlacunal, lower preocular, first subocular, and fourth supralabial. Two preoculars, upper large and convex, contacting supraocular and canthal, lower preocular thin and long; three suboculars, anterior largest and in contact with fourth supralabial; three interoculabials posterior to anterior subocular; two postoculars, upper twice as large as lower. Supralabials 13 / 13; infralabials 11 / 12; first infralabials in medial contact posterior to triangular mental; genials together resemble wings. Midbody dorsal scale rows 23–25; preventral single; ventrals 152; subcaudals 22, nondivided; eight rattle fringe scales; tail bearing three rattle segments. Ground color, in preservative, light brown with 54 dark dorsal body blotches, irregularly edged in black with a very thin light outer edge on most blotches, blotches wider than long, interspaces between blotches 1–2 scales long; smaller, vertically elongated auxiliary blotches evident laterally below the dorsal blotches, often separated from the dorsal blotches by white scales. Head marked with parallel rows of small irregular blotches across prefrontals, supraoculars, and much of the occipital area, terminated by paired occipital blotches; supralabials and infralabials ground color white, stippled with dark brown spots; distinct dark postocular stripe, uniform in width, extending from posterior of eye to above the angle of the mouth and then downwards to jawline; postocular stripe bordered by a darker band on both dorsal and ventral margins, dorsally a narrow region of light brown extends from the posterior axis of the eye along the dorsal margin of the postocular band, ventral to the postocular band is a region of white with brown stippling, supralabial 8 is nearly completely white, with moderate stippling but no postocular band, supralabials 8–13 are divided by the postocular band; two large dark brown blotches are on the borders of supralabials 5–7; scale margins of infralabials with dark brown regions making for a series of dark bands, bands do not extend beyond infralabials; gular scales stippled with no pattern; ventral scutes stippled, more heavily on posterior half of scales, with total stippling intensifying past midbody; distal 1 / 3 ventral scutes become dark brown with a lighter band along the midline, continuing to the vent. Subcaudal scales dark brown to black; eight dark brown tail bands; proximal rattle black, distal sections brown. Color in life. Color in life within the type series is only known for one paratype (MZFC 28669), shown in Fig. 8. Variation. Three of the six specimens lack a single, large anterior intercanthal. The holotype, KU 73649, has a single large intercanthal bordered anteriorly by three small, seemingly anomalous scales (Fig. 7). Similarly, UTA R- 16353 has paired anterior intercanthals separated by a small but elongated scale. The third specimen, AMNH R- 113191, has paired anterior intercanthals. The single juvenile (AMNH R- 113191) has a cream-colored proximal rattle and pale-colored tail ventrally. Variation in meristic, morphometric, and color pattern characters within the type series is listed in Table 3. Etymology. The specific epithet is a patronym honoring Jonathan A. Campbell for his many years of field research on Mexican rattlesnakes and for his decades of unwavering support to students of Mexican herpetology. Habitat and distribution. Crotalus campbelli is found in rocky open breaks within montane forest (Fig. 9) along the far western regions of the Trans-Mexican Volcanic Belt. Much of this forest is covered with remnant patches of cloud forest (Ponce-Reyes et al. 2012). This species is known from western Jalisco and the Sierra de Manantlán in southern Jalisco/northwestern Colima. A narrow low-elevation valley appears to separate the range of C. campbelli from C. armstrongi to the east (Fig. 3).