Search

We evaluated the taxonomic status of the genus Megophrys in Sumatra using molecular and morphological data. Mitochondrial phylogenetic inference and morphological data revealed two undescribed species, one in southern Sumatra—M. selatanensis sp. nov. and one in northern Sumatra—M. acehensis sp. nov. We also detected a potential cryptic species within M. parallela, but refrain from describing this lineage here due to insufficient data. Genetic variation within Sumatran Megophrys is highly structured and will require additional geographic sampling to understand the interplay between geography and genetics in Sumatran Megophrys.

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."]}

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.

Craugastor candelariensis sp. nov. Holotype. — UTA A-64253 (field ID: JAC 21885), male collected by E.N. Smith and colleagues Nof Candelaria on the road to Oaxaca; Sierra Madre del Sur, Oaxaca, Mexico, 15.94960°N, 96.47110°W, 668 m, on 21 January 2002 between 1130 and 1200 h, near stream bordering coffee plantation and secondary forest. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Paratypes (3). —MZFC-HE-35617 (formerly UTA A-64252; field ID: JAC 21873; Fig. 23), male with heavily pigmented testes, same data as holotype except collected 1.2 mi on rough road toward Pluma Hidalgo on the Candelaria – Portillo road, 15.95610°N, 96.44930°W, 1051 m, on 21 January 2002 at 1000 h in leaf litter of coffee plantation. UTA A- 66116 (Field ID: JAC 21851), male with pigmented testes collected by E.N. Smith and colleagues from San Gabriel Mixtepec, Puente de Hamaca, Oaxaca, Mexico, 16.10510°N, 97.06310°W, 710 m, on 20 January 2002 at 1520 h on forest floor. UTA A- 55247 (Field ID: ENS 9698), female with unpigmented gonads and extended oviducts collected by Karin S. Castaneda along the Carretera San Gabriel Mixtepec–Miahuatlán of the Sierra Madre del Sur, Oaxaca, Mexico, 16.160556°N, 97.00111°W, 1270–1350 m, on 15 March 1998 at 1630 h from pine forest habitat. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 18.6 mm); (2) full ossification of most skeletal elements in adults, lacking ossification only of Stage 6 (Table 3); (3) absence of posterolateral projection of frontoparietal; (4) presence of vomerine odontophores; (5) absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank with nostril– canthal–supratympanic stripe, lips colored as dorsum; (8) two postrictal tubercles; (9) gular region uniformly pale to slightly evenly peppered with melanocytes; (10) dorsal surface unicolored pale; (11) pale middorsal ridge, sometimes with few tiny spots; (12) evenly fine tubercles on dorsum; (13) body flank unicolored pale, shagreened with fine tuberculation; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and tip of snout; (16) outer tarsal ridge with 3–8 tiny and pointed tubercles on slightly raised fringe; (17) finger and toe tips lanceolate to mucronate (toes and outer two fingers); (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor candelariensis can be differentiated from C. bitonium, C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. pygmaeus, C. rubinus, and C. saltator by a larger innermetatarsal tubercle (inner and outer metatarsal tubercles are similar sizes in C. candelariensis). Craugastor candelariensis can be differentiated from C. cueyatl and C. hobartsmithi by the absence of vomerine odontophores (present in C. candelariensis). It can be differentiated from C. portilloensis by the presence of posterolateral projections of the frontoparietal (absent in C. candelariensis). Description of holotype. — Holotype small male (SVL ¼ 13.3 mm); snout rounded and short (0.5 mm naris–snout; 4% SVL); long eye–nostril distance (1.7 mm; 13% SVL); tympanum 1.2 mm (7.6% SVL); no supratympanic fold and no shoulder tubercle; finger length formula III ¼ I; single palmar tubercle; single prepollical tubercle; subarticular tubercles present on all fingers; supernumerary tubercles present on Finger III; toe length formula IV Variations in paratypes. —Body sizes (SVL) 12.4 mm (MZFC-HE-35617), 14.3 mm (UTA A-66116), 18.6 mm (UTA A-55247); eye–nostril distance 10–13% SVL (males), 9% SVL (female); tympanic ratios 7–10%. Etymology. —The name is an abbreviated allusion to the municipality of Candelaria Loxicha (near the type locality) and the Latin suffix - ensis meaning place. It is simultaneously a reference to the Latin noun candēla meaning a fire or light made of wax, given the translucent yellow appearance of several type specimens in preservative, as if someone were shining a candle through them. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Distribution. —This species is known from intermediate elevations of southern Oaxaca (668–1350 m), an area that mostly consists of Sierra Madre del Sur pine–oak forest habitat. Phylogenetics. — Craugastor candelariensis was strongly supported as monophyletic in the concatenated analysis (ML ¼ 100; BAYES ¼ 1.0; Fig. 3). In this analysis, the sister taxon of C. candelariensis was inferred to be C. polaclavus (ML ¼ 80; BAYES ¼ 0.99). We also observed this sister relationship in the nDNA-only analysis (Fig. 5); however, in the mtDNAonly analysis C. candelariensis was inferred to be the sister taxon of a clade containing C. bitonium þ C. pygmaeus (Fig. 4). In terms of genetic distances, Craugastor candelariensis is most similar to C. polaclavus and C. pygmaeus (both 6.4%; Table 4). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Remarks. —The skull of C. candelariensis is similar to C. bitonium, C. hobartsmithi, C. montanus, and C. pygmaeus, with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. Two type specimens appear white-yellowish in preservative (possibly having been saponified). This species likely co-occurs with C. pygmaeus, C. polaclavus, and C. portilloensis in southcentral Oaxaca (Figs. 6 and 8). In terms of body size and ossification level it is the smallest member of the C. mexicanus series to complete Stage 5 of our ontogenetic sequence., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 17-23, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastor polaclavus sp. nov. Holotype. — UTA A-62392 (field ID: JAC 21230; Fig. 31), female collected by E.N. Smith and colleagues in Portillo del Rayo, Distrito San Pedro Pochutla, Sierra Madre del Sur, Oaxaca, Mexico, 15.973038N, 96.997118W, 1550–1585 m, 24 September 2001. Paratypes (4). — UTA A- 55246, a recent hatchling collected by E. N. Smith and José Luís Camarillo Rangel from Río Salado, Sierra Madre del Sur, Oaxaca, Mexico, 16.1941678N, 97.09758W, 1245 m, 26 September 1997. UTA A- 66098 and MZFC-HE-35582–83, adult or subadult specimens all collected with the holotype. Referred specimen (1). — UTA A- 66097, female, same data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 14.7 mm); (2) highly reduced ossification of skeleton in adults relative to other members of series; (3) presence of posterolateral projection of the frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred with no distinctive canthal stripe, 1–2 particularly dark bars below eye; (8) one or two postrictal tubercles; (9) gular region with pale spotting; (10) dorsal surface blotched; suprascapular chevron, interorbital bar; (11) pale or as background middorsal ridge; (12) dorsum smooth with only large scattered tubercles; (13) body flank darker anteriorly, no sharp delineation of color change, smooth to shagreened; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches nostril; (16) outer tarsal ridge with 0–4 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not expanded, toe tips slightly expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor polaclavus can be differentiated from C. candelariensis, C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by the presence of vomerine odontophores (absent in C. polaclavus). It can be differentiated from C. bitonium, and C. cueyatl by the condition of adpressed leg where the heel does not reach the nostril (reaches nostril in C. polaclavus). It can be differentiated from C. hobartsmithi and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. polaclavus). It can be differentiated from C. portilloensis by metatarsal tubercles of equal size (unequal in C. polaclavus). It can be differentiated from C. rubinus by relative finger lengths of IV ¼ II (IV> II in C. polaclavus). Description of holotype. — Holotype small female (SVL ¼ 14.7 mm); snout rounded and short (0.8 mm naris–snout; 5% SVL); short eye–nostril distance (1.4 mm; 7% SVL); tympanum 1.4 mm (10% SVL); mild supratympanic fold terminating in small shoulder tubercle; finger length formula III ¼ II Variations in paratypes. —Body sizes in SVL 8.2 mm (UTA A-55246), 12.3 mm (UTA A-66098), 11.6 mm (MZFC-HE-35582), 13.5 mm (MZFC-HE-35583); eye–nostril distance 11–13% SVL; tympanic ratios 7–8%. Distribution. —Intermediate elevations in the foothills of the Sierra Madre del Sur in Oaxaca 1245–1585 m (Fig. 7). These habitats consist of mixed tropical dry and temperate sierra forests. Etymology. —The specific epithet is a combination of the Latin pola meaning small and clavus meaning wart. The name is an allusion to the small size and rugose appearance of several individuals in the type series. Phylogenetics. — Craugastor polaclavus was inferred to be the sister taxon of C. candelariensis with strong support in the concatenated analysis (90 ML; 0.99 BAYES; Fig. 3) and nDNA-only analysis (ML> 90, BAYES> 0.90; Fig. 5). The placement of C. polaclavus was less certain in the mtDNA, where it was found to be the sister taxon of a clade containing C. cf. hobartsmithi þ C. rubinus (ML ¼ 51, BAYES ¼ 0.74; Fig. 4). In terms of genetic distances (Table 4), C. polaclavus was most similar to C. portilloensis (5.8%), followed by similarity to C. bitonium (5.9%). Remarks. —The skull of C. polaclavus is similar to C. mexicanus, C. omiltemanus, and C. saltator, with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. Specimens were dissected; three (UTA A-62392, 66098, and MZFC-HE-35582) seem to be subadult females with unpigmented ovaries and thin undeveloped oviducts, UTA A-66097 is an adult female with a thickened oviduct and also unpigmented ovaries containing yolked eggs, UTA A-66098 adult and MZFC-HE-35582 subadult are males with pigmented testes (smaller on second specimen), and the hatchling (8.2 mm SVL) was not dissected. This species likely co-occurs with C. candelariensis, C. portilloensis, and C. pygmaeus in southcentral Oaxaca (Figs. 6 and 8). It was collected in sympatry with C. portilloensis at Portillo del Rayo, Oaxaca, Mexico (Fig. 8). The hatchling paratype specimen (UTA A-55246) had no evidence of skeletal ossification (Fig. 11).

Craugastor pygmaeus (Taylor 1936) Eleutherodactylus pygmaeus Taylor 1936:352. Holotype female (UIMNH 16125) from ‘‘one mile north of Rodriguez Clara, Veracrux, Mexico.’’ [Examined]. Microbatrachylus albolabris Taylor 1940:502. Holotype female (FMNH 100071) from ‘‘two miles west of Córdoba, Veracruz, Mexico.’’ [Examined]. Microbatrachylus pygmaeus (Taylor): Taylor 1940:500. Microbatrachylus minimus Taylor 1940:507. Holotype male (FMNH 100323) from ‘‘Agua del Obispo, Guerrero, Mexico.’’ [Examined]. Microbatrachylus imitator Taylor 1942:70. Holotype female (USNM 115508) from ‘‘La Esperanza, Chiapas, Mexico.’’ [Examined]. Craugastor pygmaeus (Taylor): Crawford and Smith 2005:536. Diagnosis. —Based on 37 specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) small adult size (mean SVL ¼ 13.5 mm [SD ¼ 1.77], n ¼ 29); (2) full ossification of skeletal elements in adults; (3) lack of posterolateral projection of frontoparietal; (4) lack of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred canthus and jaws (rarely dark and blotched), no canthal stripe; (8) one (or two fused) postrictal tubercles; (9) gular region pigmentation present or absent; (10) dorsal surface two-toned usually with dark suprascapular ^^shape, or striped and with pale middorsal stripe; (11) variable middorsal ridge; (12) dorsum smooth or with only some large scattered tubercles; (13) body flank barred darker anteriorly, slightly shagreened to smooth; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and slightly beyond tip of snout; (16) outer tarsal ridge smooth, no raised fringe; (17) finger and toe pads round, expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor pygmaeus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. pygmaeus). It can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. rubinus, and C. saltator by the presence of a posterolateral projection of the frontoparietal (absent in C. pygmaeus ; Fig. 26C). Craugastor pygmaeus is most similar to C. bitonium (in morphology, osteology, and genetic distance), but may be differentiated from this taxon by the condition of the outer tarsal ridge, which is smooth in C. pygmaeus versus 1–6 small tubercles in C. bitonium. Description. —In previous literature, described as smallbodied (‘‘diminutive’’) and short-limbed, with unequal inner and outer metatarsal tubercle sizes; distinct subarticular tubercles; barely visible supernumerary tubercles; no vomerine odontophores (Taylor 1936); rounded canthus; two palmar tubercles (Taylor 1942). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Holotype (UIMNH 16125) ~ 18 mm SVL. Among select specimens that we examined, we observed short, rounded snout (naris–snout ¼ 0.78 mm, SD ¼ 0.14, n ¼ 29; 5.8% SVL); long eye–nostril distance (1.19 mm, SD ¼ 0.24, n ¼ 29; 8.9% SVL); some specimens with single palmar tubercle; relative finger lengths III> IV> II ¼ I; relative toe lengths IV Distribution. —Widely distributed throughout lowland to intermediate localities in the states of Chiapas, Puebla, Oaxaca, and Veracruz (also possibly Tabasco, Mexico, and western Guatemala) from near sea level to 2000 m (Fig. 6). Canseco Márquez and Gutiérrez Mayén (2010) report C. pygmaeus occurs in the forests adjacent to the Tehuacán– Cuicatlán Valley in Puebla and Oaxaca. The western range edge of C. pygmaeus is uncertain; we examined one specimen (UTA A-66131) a male with pigmented testes, from San Vicente de Benitez (17.290618N, 100.279558W, 951 m), Guerrero, collected 17 June 2004, that appears referable to C. pygmaeus. Diet. —One male specimen (UTA A-64263, determined by the presence of pigmented testes), was found to contain ants (Formicidae) in its stomach. Phylogenetics. — Craugastor pygmaeus was inferred to be the sister taxon of C. bitonium with high support in the concatenated analyses (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). This sister relationship was also recovered in both mtDNA and nDNA analyses, although with lower support in the nDNAonly analyses (ML ¼ 54, BAYES ¼ 0.67; Figs. 4 and 5). Craugastor pygmaeus is separated from C. bitonium by a P - distance of 4.7% (Table 4). Remarks. —The skull of C. pygmaeus is similar to C. hobartsmithi and C. montanus, with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. Despite how common this species is in most museum collections, we know very little about its natural history, including reproductive behavior, call, and diet. This species likely co-occurs with C. candelariensis, C. polaclavus and C. portilloensis in southcentral Oaxaca and (possibly) C. bitonium in central Guerrero. It may overlap with (1) C. montanus in Chiapas, (2) C. mexicanus at intermediate elevations of the Sierra Madre del Sur and Sierra Madre Oriental (Fig. 7), and (3) C. hobartsmithi in central and western Guerrero. Males likely have larger tympana than do females (Figs. 13 and 33)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 34-35, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1936 [1937]. New species of amphibia from Mexico. Transactions of the Kansas Academy of Science 39: 349 - 363.","Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571.","Taylor, E. H. 1942. New tailless Amphibia from Mexico. University of Kansas Science Bulletin 28: 67 - 89.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Canseco Marquez, L., and M. G. Gutierrez Mayen. 2010. Anfibios y Reptiles del Valle Tehuacan-Cuicatlan. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Mexico."]}

Craugastor rubinus sp. nov. Holotype. — UTA A-62345 (field ID: JAC 30720; Fig. 34A), male collected by J.W. Streicher, C.L. Cox, C.M. Sheehy, III, R.U. Tovar, and M.J. Ingrasci on the road between Talpa de Allende and El Cuale, Jalisco, Mexico, 20.377078N, 105.047938W, 1771 m, 8 July 2009. Paratypes (2). — UTA A- 62347 (Fig. 34 C) and MZFC-HE-35616 (formerly UTA A- 62346; Fig. 34 B), same collection data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size; (2) reduced ossification of the skeleton in adults relative to other members of series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred, with snout–nostril–canthal–supratympanic stripe, sometimes broken canthaly and postocularly; (8) one postrictal tubercle; (9) gular region with pale spotting; (10) dorsal surface blotched or unicolored pale; diffuse interorbital bar, small suprascapular spots, sometimes two dark rump spots; (11) middorsal ridge present; (12) dorsum smooth with no tubercles; (13) body flank dark supratympanic stripe extending toward lower midflank, broken, paler toward groin, smooth to finely shagreened; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches nostril; (16) outer tarsal ridge with 0–4 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not or just barely expanded, toe tips slightly lanceolate and barely expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor rubinus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. rubinus). It can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, and C. saltator by the presence of vomerine odontophores (absent in C. rubinus). It can be differentiated from C. bitonium and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. rubinus). Description of holotype. — Holotype small male (SVL ¼ 12.6 mm); snout rounded and short (0.8 mm naris–snout; 7% SVL); short eye–nostril distance (1.3 mm; 10% SVL); tympanum 1.2 mm (9.1% SVL); no supratympanic fold or shoulder tubercle; finger length formula III C. mexicanus series (Fig. 35); grey in preservative; many bands present on arms and legs. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Variations in paratypes. —Body sizes (SVL) 10.8 mm (MZFC-HE-35616), 11.5 mm (UTA A-62347); eye–nostril distance both 10% SVL; tympanic ratios both 9%; dorsal color pattern mottled (MZFC-HE-35616; Fig. 34B) or with dark stripe extending from snout to flank (UTA A-62347, Fig. 34C). Distribution. —Known only from the type locality in Jalisco (Talpa de Allende), habitat of pine–oak forest at the southern extent of the Sierra Madre Occidental. Etymology. —The specific epithet is derived from the Latin rubinus, which means ruby. This name is inspired by the garnet mines found near the type locality of Talpa de Allende in the Sierra Madre Occidental of Jalisco. Phylogenetics. — Craugastor rubinus was strongly supported as monophyletic in the concatenated analysis (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). The new species is also strongly supported as the sister taxon of C. cf. hobartsmithi (ML ¼ 100; BAYES ¼ 1.0). Craugastor rubinus is separated from C. cf. hobartsmithi by a P -distance of 3.4% (Table 4). Remarks. —The skull of C. rubinus is similar to that of C. hobartsmithi, C. montanus, and C. pygmaeus with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. The type series was collected from the leaf litter surrounding a mountain stream. The male holotype has slightly pigmented testes. Afemale paratype, UTA A-62345, has large, pigmented ovaries. This species may co-occur with C. hobartsmithi in Jalisco. Despite the small genetic distances separating C. rubinus and C. cf. hobartsmithi , there are many skeletal and morphological differences between these two species, including relative metatarsal tubercle sizes, condition of the posterolateral projection of frontoparietal, and relative snout length (Table 6)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 35-37, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastor portilloensis sp. nov. Holotype. — UTA A-62393 (field ID: JAC 21431; Fig. 32), subadult female collected by E.N. Smith and colleagues in Portillo del Rayo, Distrito San Pedro Pochutla, Sierra Madre del Sur, Oaxaca, Mexico, 15.9794448N, 96.5166678W, 1550 m, 1 October 2001. Paratypes (4). —MZFC-HE-35580 and UTA A- 66095, juvenile males. MZFC-HE- 35581 subadult female, and UTA A- 66096 subadult male. All collected at the type locality between 1550 and 1585 mon 24 September 2001 by E.N. Smith and colleagues. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 11.4 mm); (2) reduced ossification of skeleton in adults relative to other members of the series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank darker than dorsum, faintly barred lips, snout–nostril– canthal–supratympanic stripe; (8) one or two postrictal tubercles; (9) gular region with pale spotting; (10) dorsal surface unicolored pale, diffuse interorbital bar, sometimes with two dark rump spots; (11) with or without a middorsal ridge; (12) dorsum smooth or with scattered fine tubercles; (13) body flank has dark supratympanic stripe extending toward lower mid-flank, smooth, very few small tubercles; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and slightly past snout; (16) outer tarsal ridge smooth or with 1–3 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not expanded, toe tips slightly expanded; (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor portilloensis can be differentiated from C. candelariensis, C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by the presence of vomerine odontophores (absent in C. portilloensis). It can be differentiated from C. bitonium, C. hobartsmithi, and C. pygameus by the absence of a posterolateral projection of the frontoparietal (present in C. portilloensis). It can be further differentiated from C. pygmaeus by the absence of a canthal mask (present in C. portilloensis). It can be differentiated from C. cueyatl and C. polaclavus by metatarsal tubercles of unequal size (equal in C. portilloensis). It can be differentiated from C. rubinus by relative finger lengths of IV ¼ II (IV> II in C. portilloensis). Description of holotype. — Holotype small female (SVL ¼ 11.4 mm); snout rounded and short (0.8 mm naris–snout; 6% SVL); short eye–nostril distance (1.1 mm; 9.8% SVL); tympanum 0.8 mm (6.8% SVL); no supratympanic fold; small shoulder tubercle; finger length formula III Variations in paratypes. —Body sizes in SVL 8.7 mm (MZFC-HE-35580), 8.4 mm (UTA A-66095), 12.1 mm (MZFC-HE-35581), 12.6 mm (UTA A-66096); eye–nostril distance 10–11% SVL; tympanic ratios 7%. Etymology. —The name is an abbreviated allusion to the type locality near the town of Portillo del Rayo and the Latin suffix - ensis meaning place. Distribution. —Intermediate elevations in the foothills of the Sierra Madre del Sur in Oaxaca 1550–1585 m (Fig. 8). These habitats consist of mixed tropical dry and temperate sierra forests. Phylogenetics. —In the concatenated analysis, Craugastor portilloensis is the sister taxon to a clade of small-bodied species (C. bitonium, C. candelariensis, C. cueyatl, C. hobartsmithi, C. polaclavus, C. pygmaeus, and C. rubinus), but with low branch support (ML ¼ 53, BAYES ¼ 66; Fig. 3). It occupies the same phylogenetic placement in the mtDNA-only data set (Fig. 4), but in the nDNA-only data set is the sister taxon of a clade including (C. bitonium, C. cueyatl, C. hobartsmithi, C. pygmaeus, and C. rubinus) with moderate support in the Bayesian analysis (ML ¼ 43, BAYES ¼ 0.78; Fig. 5). In terms of genetic distances (Table 4), C. portilloensis was most similar to C. montanus (5.7%), followed by similarity to C. polaclavus (5.8%). Remarks. —The skull of C. portilloensis is similar to C. mexicanus, C. omiltemanus, and C. saltator, with a more anteriorly placed anterior suture of frontoparietal and prootic than in other species. We dissected all specimens of C. portilloensis; males possess pigmented testes and females unpigmented ovaries. This species likely co-occurs with C. candelariensis, C. polaclavus, and C. pygmaeus in southcentral Oaxaca (Figs. 6 and 8). It was collected in sympatry with C. polaclavus at the type locality of Portillo del Rayo, Oaxaca, Mexico (Fig. 8).

Craugastor hobartsmithi (Taylor 1936) Eleutherodactylus hobartsmithi Taylor 1936:355. Holotype male (FMNH 100114) from ‘‘Uruapan, Michoacán, Mexico.’’ [Examined]. Microbatrachylus hobartsmithi (Taylor): Taylor 1940:501. Craugastor hobartsmithi (Taylor): Crawford and Smith 2005:536. Microbatrachylus pygmaeus Duellman 1961:34. [Misidentification]. Craugastor pygmaeus Ahumada-Carrillo et al. 2013:1338. [Misidentification]. Diagnosis. —Based on six specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) small adult sizes (males 11.3–15.2 mm, females ~ 16.7 mm; Table 5); (2) full ossification of most skeletal elements in adults, lacking ossification beyond Stage 4 (Table 3); (3) absence of vomerine odontophores; (4) absence of posterolateral projection of frontoparietal; (5) presence of a row of 4–6 rounded tubercles along outer edge, and 1–3 in mid upper eyelid; (6) supratympanic fold poorly developed; (7) face flank barred with or without distinctive canthal stripe; (8) one or two postrictal tubercles; (9) gular region peppered with melanocytes; (10) dorsal surfaces finely blotched, usually with dark interorbital bar and suprascapular ^-shape, some individuals with pale dorsal color and four stripes, paravertebral and lateral, originating at corners of eyes and ending above groin (lateral more prominent); (11) middorsal ridge (dark or background color); (12) mostly smooth dorsum or with just fine tubercles or folds toward back; (11) body flank darker anteriorly, around axilla, slightly tubercular; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between anterior corner of eye and snout; (16) outer tarsal ridge with 4–6 rounded to slightly pointed tubercles, no raised fringe; (17) finger and toe tips round, finger tips slightly expanded, toe tips expanded; (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor hobartsmithi can be differentiated from C. bitonium, C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. pygmaeus, and C. rubinus by an inner metatarsal tubercle that is twice the size of the outer (these are similar sizes in C. hobartsmithi). It can be differentiated from C. candelariensis and C. saltator by the presence of vomerine odontophores (absent in C. hobartsmithi). It can be differentiated from C. cueyatl and C. portilloensis by the presence of posterolateral projections of the frontoparietal (absent in C. hobartsmithi). Description. —In previous literature, C. hobartsmithi has been described as small-bodied with pigmented gonads (Taylor 1936, 1940); presence of tubercles on the tarsus (Duellman 1961); two palmar tubercles (Lynch 1965); tarsus bearing a row of tubercles along its outer edge (Lynch 1970). Holotype (FMNH 100114) small male (13.5 mm); snout rounded and short (0.9 mm naris–snout; 6% SVL); short eye–nostril distance (1.18 mm; 8.7% SVL); tympanum 1.9 mm (14% SVL). We further examined two specimens of C. cf. hobartsmithi from coastal Michoacán (UTA A-66133–34; Fig. 25B and C) and noted the following characteristics: supratympanic fold terminating in two postrictal tubercles; finger length formula III ¼ II Distribution. — Craugastor hobartsmithi occurs in the pine–oak forest of Michoacán. Craugastor cf. hobartsmithi occurs throughout western Mexico in low to intermediate habitats of Jalisco, Nayarit, Michoacán, Guerrero, and Sinaloa (Fig. 8; Hardy and McDiarmid 1969). FloresCobarrubias et al. (2012) reported C. hobartsmithi from Hostotipaquillo, Jalisco. García and Ceballos (1994) reported C. hobartsmithi from coastal Jalisco. The records of C. pygmaeus reported in Duellman (1961) and AhumadaCarrillo et al. (2013) are all likely C. hobartsmithi or C. cf. hobartsmithi because our molecular results indicate that C. pygmaeus does not occur west of Guerrero. Similarly, many iNaturalist (https://www.inautralist.org, accessed June 2019) accounts of C. cf. hobartsmithi are listed under C. pygmaeus —these accounts also suggest that C. cf. hobartsmithi is much more widely distributed in western Mexico than museum collections indicate. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Phylogenetics. — Craugastor cf. hobartsmithi was found to be the sister taxon of C. rubinus with strong support (ML ¼ 100; BAYES ¼ 1.0) in both the concatenated and separate mtDNA and nDNA analyses (Figs. 3, 4, and 5). The pairwise P -distances between C. cf. hobartsmithi and C. rubinus is 3.4% (Table 4); this is the smallest genetic distance between any species of the C. mexicanus series, suggesting recent divergence. Remarks. —The skull of C. hobartsmithi is similar to C. montanus and C. pygmaeus, with more posteriorly placed anterior suture of frontoparietal and prootic than in other species. We tentatively referred several museum collections to C. hobartsmithi (Fig. 8) as C. cf. hobartsmithi , but these should be further examined. The specimens of C. pygmaeus reported by Duellman (1961) from Arteaga, Michoacán, are referred to C. cf. hobartsmithi because we examined several C. pygmaeus from Oaxaca with tubercles on the outer edge of the tarsus rendering Duellman’s (1961) apomorphic character for C. hobartsmithi unreliable. Craugastor hobartsmithi may co-occur with C. pygmaeus in southcentral Guerrero (Figs. 6 and 8). The tissues of C. cf. hobartsmithi used in our phylogenetic analysis originated from Colima. Although we lack a voucher specimen for the tissue, the collector of the tissue (J. Reyes-Velasco) provided us with photographs of C. cf. hobartsmithi from Montitlan, near where the tissue was collected (Fig. 25E and F). One female specimen of C. hobartsmithi from near Uruapan (UMMZ 94230) had several intradermal trombiculid mites on its venter (Fig. 15D)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 25-27, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1936 [1937]. New species of amphibia from Mexico. Transactions of the Kansas Academy of Science 39: 349 - 363.","Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Duellman, W. E. 1961. The amphibians of Michoacan, Mexico. Miscellaneous Publications of the University of Kansas Museum of Natural History 15: 1 - 148.","Ahumada-Carrillo, I. T., J. C. Arenas-Monroy, F. A. Fernandez-Nava, and O. Vazquez-Huizar. 2013. New distributional records for the pygmy robber frog Craugastor pygmaeus (Terrarana: Craugastoridae) in western Mexico. Revista Mexicana de Biodiversidad 84: 1338 - 1342.","Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA.","Lynch, J. D. 1970. Taxonomic notes on some Mexican frogs (Eleutherodactylus: Leptodactylidae). Herpetologica 26: 172 - 180.","Hardy, L., and R. McDiarmid. 1969. The amphibians and reptiles of Sinaloa, Mexico. University of Kansas Publications, Museum of Natural History 18: 39 - 252.","Garcia, A., and G. Ceballos. 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. [In Spanish.]"]}

Craugastor bitonium sp. nov. Holotype. — UTA A-64254 (field ID: JAC 22117; Fig. 22A), adult female from road between Yerba Santa and Yextla (HWY 196), Guerrero, Mexico, 17.52666°N, 99.9579°W, 2071 m, collected by J.A. Campbell and colleagues on 10 June 2002. Paratypes (5). —MZFC-HE-35600–01 and UTA A- 66117– 18 adult females, and UTA A-66119 adult male (Fig. 22B–D), all same collection data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 16.7 mm); (2) full ossification of skeletal elements in adults; (3) absence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred or with supralabial pale stripe, and with or without dark canthal stripe; (8) single postrictal tubercle; (9) gular region peppered with melanocytes; (10) dorsal surface two-toned, usually with a dark suprascapular ^ shape, or almost unicolored; (11) pale or ground color middorsal ridge; (12) scattered fine tubercles on dorsum; (13) body flank barred darker anteriorly, slightly shagreened to smooth; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches middle of eye to slightly beyond snout; (16) outer tarsal ridge with 1–6 small mostly round tubercles, no raised fringe; (17) finger and toe tips round, slightly lanceolated, slightly expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor bitonium can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by their large adult body sizes of SVL> 20 mm (C. bitonium), the presence of vomerine odontophores (absent in C. bitonium), and the presence of three palmar tubercles (one palmar tubercle in C. bitonium). Craugastor bitonium can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by the presence of metatarsal tubercles of similar sizes (different sizes in C. bitonium). Craugastor bitonium can be differentiated from C. rubinus by the presence of posterolateral projections of the frontoparietal (absent in C. bitonium). Craugastor bitonium can be differentiated from C. polaclavus by its shorter eye–nostril distance with an eye–nostril distance 9–10% SVL in C. bitonium and 10–13% SVL in C. polaclavus. Craugastor bitonium is most similar to C. pygmaeus (in morphology, osteology, and genetic distance), but may be differentiated from this taxon by the condition of the outer tarsal ridge; 1–6 small tubercles in C. bitonium versus no tubercles (¼ smooth) in C. pygmaeus. Holotype description. — Holotype small female with unpigmented developing ova (SVL ¼ 15.8 mm); snout rounded and short (0.9 mm naris–snout; 6% SVL); short eye–nostril distance (1.42 mm; 9% SVL); tympanum 1.4 mm (8.9% SVL); small supratympanic fold terminating in shoulder tubercle; finger length formula III Variations in paratypes. —Body sizes (SVL) 15.8 mm (MZFC-HE-35600), 15.2 mm (MZFC-HE-35601), 16.9 mm (UTA A- 66117), 16.7 mm (UTA A- 66118), 12.3 mm (UTA A- 66119); eye–nostril distance 9–10% SVL; tympanic ratios 7– 9%; dorsal color patterns variable, often with two distinctive patches of differing ground coloration ranging from orange to tan. Etymology. —The specific epithet is a combination of the Latin prefix bi- meaning two and tonium meaning tone. It is a reference to the two distinctive patches of color found on the holotype and several paratypes that create the appearance of a ‘‘two-tone’’ dorsal coloration. Distribution. —This species is known only from the Sierra Madre del Sur of central Guerrero (~ 2071 m). The closest named places to the type locality are Izotepec to the north and Los Bajos to the southwest. The habitat at the type locality is montane pine–oak forest. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Diet. —CT-scan of the holotype revealed the presence of a small millipede (Diplopoda) in the stomach. We also noted the presence of a small red ant (Formicidae) in the mouth of the holotype. Phylogenetics. — Craugastor bitonium was inferred to be the sister taxon of C. pygmaeus, with high support in the concatenated analyses (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). This sister relationship was also recovered in both mtDNA and nDNA analyses, although with lower support in the nDNAonly analyses (ML ¼ 54, BAYES ¼ 0.67; Figs. 4 and 5). Craugastor bitonium is separated from C. pygmaeus by a P - distance of 4.7% (Table 4). Remarks. —The skull of C. bitonium is similar to C. hobartsmithi, C. montanus, and C. pygmaeus. Craugastor bitonium displays a developmental pattern similar to C. pygmaeus (with high levels of ossification at small sizes), suggesting small adult body sizes (Fig. 11). This species likely co-occurs with C. pygmaeus, C. omiltemanus, and C. saltator in central Guerrero (Figs. 6 and 8). The specimen UTA A-66132 is referred with some hesitation because it was collected from a lower elevation than the type locality and has a Finger Ilength nearing C. pygmaeus (which also occurs in Guerrero). Six of the specimens were adult females containing unpigmented ovaries with yolked eggs and thick oviducts, the seventh (UTA A-66119; Fig. 22C, far left) was an adult male with pigmented testes., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 13-16, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastor rubinus sp. nov. Holotype. — UTA A-62345 (field ID: JAC 30720; Fig. 34A), male collected by J.W. Streicher, C.L. Cox, C.M. Sheehy, III, R.U. Tovar, and M.J. Ingrasci on the road between Talpa de Allende and El Cuale, Jalisco, Mexico, 20.377078N, 105.047938W, 1771 m, 8 July 2009. Paratypes (2). — UTA A- 62347 (Fig. 34 C) and MZFC-HE-35616 (formerly UTA A- 62346; Fig. 34 B), same collection data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size; (2) reduced ossification of the skeleton in adults relative to other members of series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred, with snout–nostril–canthal–supratympanic stripe, sometimes broken canthaly and postocularly; (8) one postrictal tubercle; (9) gular region with pale spotting; (10) dorsal surface blotched or unicolored pale; diffuse interorbital bar, small suprascapular spots, sometimes two dark rump spots; (11) middorsal ridge present; (12) dorsum smooth with no tubercles; (13) body flank dark supratympanic stripe extending toward lower midflank, broken, paler toward groin, smooth to finely shagreened; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches nostril; (16) outer tarsal ridge with 0–4 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not or just barely expanded, toe tips slightly lanceolate and barely expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor rubinus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. rubinus). It can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, and C. saltator by the presence of vomerine odontophores (absent in C. rubinus). It can be differentiated from C. bitonium and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. rubinus). Description of holotype. — Holotype small male (SVL ¼ 12.6 mm); snout rounded and short (0.8 mm naris–snout; 7% SVL); short eye–nostril distance (1.3 mm; 10% SVL); tympanum 1.2 mm (9.1% SVL); no supratympanic fold or shoulder tubercle; finger length formula III C. mexicanus series (Fig. 35); grey in preservative; many bands present on arms and legs. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Variations in paratypes. —Body sizes (SVL) 10.8 mm (MZFC-HE-35616), 11.5 mm (UTA A-62347); eye–nostril distance both 10% SVL; tympanic ratios both 9%; dorsal color pattern mottled (MZFC-HE-35616; Fig. 34B) or with dark stripe extending from snout to flank (UTA A-62347, Fig. 34C). Distribution. —Known only from the type locality in Jalisco (Talpa de Allende), habitat of pine–oak forest at the southern extent of the Sierra Madre Occidental. Etymology. —The specific epithet is derived from the Latin rubinus, which means ruby. This name is inspired by the garnet mines found near the type locality of Talpa de Allende in the Sierra Madre Occidental of Jalisco. Phylogenetics. — Craugastor rubinus was strongly supported as monophyletic in the concatenated analysis (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). The new species is also strongly supported as the sister taxon of C. cf. hobartsmithi (ML ¼ 100; BAYES ¼ 1.0). Craugastor rubinus is separated from C. cf. hobartsmithi by a P -distance of 3.4% (Table 4). Remarks. —The skull of C. rubinus is similar to that of C. hobartsmithi, C. montanus, and C. pygmaeus with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. The type series was collected from the leaf litter surrounding a mountain stream. The male holotype has slightly pigmented testes. Afemale paratype, UTA A-62345, has large, pigmented ovaries. This species may co-occur with C. hobartsmithi in Jalisco. Despite the small genetic distances separating C. rubinus and C. cf. hobartsmithi , there are many skeletal and morphological differences between these two species, including relative metatarsal tubercle sizes, condition of the posterolateral projection of frontoparietal, and relative snout length (Table 6).

Craugastor montanus (Taylor 1942) Microbatrachylus montanus Taylor 1942:67. Holotype female (USNM 115507) from ‘‘Mount Ovando, Chiapas, Mexico.’’ [Examined]. Eleutherodactylus sartori Lynch 1965:10. [Replacement name]. Craugastor montanus (Taylor): Crawford and Smith 2005:536. Diagnosis. —Based on holotype (Fig. 1E). Aspecies of Craugastor distinguished by the following combination of characters: (1) moderate adult size (holotype, SVL ¼ 24.5 mm); (2) ossification of most of skeleton in adults; (3) presence of posterolateral projection of frontoparietal; (4) presence of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold moderate to poorly developed; (7) face flank barred with or without canthal stripe, 1–2 particularly dark bars below eye; (8) one (or two fused) postrictal tubercles; (9) gular region with pale spotting; (10) dorsal surface blotched or unicolored pale; diffuse interorbital bar, small suprascapular spots; sometimes with two dark rump spots (11) middorsal ridge present; (12) dorsum smooth with only few fine tubercles; (13) body flank darker anteriorly (post axillary), slightly shagreened to smooth; (14) inguinal gland present and axillary gland present in adults; (15) when leg adpressed to body, heel reaches snout tip or beyond; (16) outer tarsal ridge with 0–2 small and round tubercles close to heel, no raised fringe; (17) finger and toe tips round, finger tips slightly or not expanded, toe tips expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor montanus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. montanus). It can be differentiated from C. bitonium and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. montanus). It can be differentiated from C. polaclavus and C. rubinus by the absence of vomerine odontophores (present in C. montanus). It can be differentiated from C. mexicanus, C. omiltemanus, and C. saltator by the general shape of its skull (Fig. 12). It can be differentiated from C. mexicanus by the condition of supratympanic folds in adults; moderate to poorly developed in C. montanus versus developed in C. mexicanus. It can be differentiated from C. omiltemanus by ventral skin texture in life; smooth to granular in C. montanus versus areolate in C. omiltemanus. It can be differentiated from C. saltator by relative leg length; crus 53–59% SVL in C. montanus versus 62–73% SVL in C. saltator. Description. —In previous literature, described as moderately sized (males average 16.2 mm SVL, females average 24.0 mm SVL); Finger Ishorter than Finger II; three palmar tubercles; testes black; inner metatarsal tubercle larger than outer metatarsaltubercle (Lynch 2000). Lynch (2000:133) redescribed C. montanus (as E. sartori) owing to ‘‘errors in Taylor’s (1942) original description that were repeated by Lynch (1965, 1970)’’. Distribution. —This species is known from intermediate to high elevations (~ 2000 m) of the Sierra Madre de Chiapas in the state of Chiapas, Mexico (Lynch 2000), and adjacent regions of the Department of San Marcos, Guatemala (Crawford and Smith 2005). This region contains a complex mixture of dry forests, mixed forests, cloud forests, and pine– oak forests. Phylogenetics. —In the concatenated analysis, C. montanus was recovered as the sister taxon of all other members of the C. mexicanus series (ML ¼ 66, BAYES ¼ 0.99; Fig. 3). It also had this placement in the mtDNA-only analysis (Fig. 4), but not in the nDNA analysis where it was found with weak support to be the sister taxon of a clade containing all taxa except C. mexicanus and C. omiltemanus (ML ¼ 33, BAYES ¼ 0.73; Fig. 5). This differs from the phylogenetic placement of C. montanus (as E. sartori) in the nDNA-only analysis of Crawford and Smith (2005). In terms of genetic distances (Table 4), C. montanus was most similar to C. polaclavus (5.7%) followed by similarity with C. mexicanus (6.1%). Remarks. —The skull of C. montanus is similar to C. hobartsmithi and C. pygmaeus, with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. The skull of C. montanus was also described by Lynch (2000) as E. sartori. Lynch (1965) created the neonym, Eleutherodactylus sartori, because Eleutherodactylus montanus was preoccupied by a West Indian species. This is the most southernly distributed species in the C. mexicanus series. Craugastor montanus likely co-occurs with C. pygmaeus. The type locality of C. greggi (Bumzahem 1955) is Volcan Tajumulco in San Marcos, Guatemala near where we sampled C. montanus for our molecular analysis (Fig. 6). Although C. greggi was placed in the C. laticeps series of Hedges et al. (2008), it is allied to the C. mexicanus series by having Finger Ishorter than Finger II. This affinity was noted in the original description: ‘‘ Eleutherodactylus greggi seems to agree most closely with the member of the Eleutherodactylus mexicanus group...’’ (Bumzahem 1955:119). However, C. greggi was differentiated from C. montanus (¼ E. sartori) by Lynch (2000) on the basis of fusion between the last presacral vertebrae and sacrum (not fused in C montanus). Nonetheless, our preliminary examinations of the holotype of C. greggi (Fig. 28) and collections from the Sierra de Chiapas suggest that future research is needed to confirm C. greggi can be differentiated from C. montanus because the taxa are united by multiple characters including (but not limited to) adult body size, presence of vomerine odonotophores, presence of posterolateral projection of frontoparietal, gular region with pale spotting, finger lengths, toe lengths, unequal sizes of the inner and outer metatarsal tubercles, and geographic distribution., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on page 29, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1942. New tailless Amphibia from Mexico. University of Kansas Science Bulletin 28: 67 - 89.","Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Lynch, J. D. 1970. Taxonomic notes on some Mexican frogs (Eleutherodactylus: Leptodactylidae). Herpetologica 26: 172 - 180.","Bumzahem, C. B. 1955. A New species of frog of the genus Eleutherodactylus from Guatemala. Copeia 1955: 118 - 119.","Hedges, S. B., W. E. Duellman, and M. P. Heinicke. 2008. New World Direct-Developing Frogs (Anura: Terrarana): Molecular Phylogeny, Classification, Biogeography, and Conservation. Magnolia Press, New Zealand."]}

Craugastor candelariensis sp. nov. Holotype. — UTA A-64253 (field ID: JAC 21885), male collected by E.N. Smith and colleagues Nof Candelaria on the road to Oaxaca; Sierra Madre del Sur, Oaxaca, Mexico, 15.94960°N, 96.47110°W, 668 m, on 21 January 2002 between 1130 and 1200 h, near stream bordering coffee plantation and secondary forest. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Paratypes (3). —MZFC-HE-35617 (formerly UTA A-64252; field ID: JAC 21873; Fig. 23), male with heavily pigmented testes, same data as holotype except collected 1.2 mi on rough road toward Pluma Hidalgo on the Candelaria – Portillo road, 15.95610°N, 96.44930°W, 1051 m, on 21 January 2002 at 1000 h in leaf litter of coffee plantation. UTA A- 66116 (Field ID: JAC 21851), male with pigmented testes collected by E.N. Smith and colleagues from San Gabriel Mixtepec, Puente de Hamaca, Oaxaca, Mexico, 16.10510°N, 97.06310°W, 710 m, on 20 January 2002 at 1520 h on forest floor. UTA A- 55247 (Field ID: ENS 9698), female with unpigmented gonads and extended oviducts collected by Karin S. Castaneda along the Carretera San Gabriel Mixtepec–Miahuatlán of the Sierra Madre del Sur, Oaxaca, Mexico, 16.160556°N, 97.00111°W, 1270–1350 m, on 15 March 1998 at 1630 h from pine forest habitat. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 18.6 mm); (2) full ossification of most skeletal elements in adults, lacking ossification only of Stage 6 (Table 3); (3) absence of posterolateral projection of frontoparietal; (4) presence of vomerine odontophores; (5) absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank with nostril– canthal–supratympanic stripe, lips colored as dorsum; (8) two postrictal tubercles; (9) gular region uniformly pale to slightly evenly peppered with melanocytes; (10) dorsal surface unicolored pale; (11) pale middorsal ridge, sometimes with few tiny spots; (12) evenly fine tubercles on dorsum; (13) body flank unicolored pale, shagreened with fine tuberculation; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and tip of snout; (16) outer tarsal ridge with 3–8 tiny and pointed tubercles on slightly raised fringe; (17) finger and toe tips lanceolate to mucronate (toes and outer two fingers); (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor candelariensis can be differentiated from C. bitonium, C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. pygmaeus, C. rubinus, and C. saltator by a larger innermetatarsal tubercle (inner and outer metatarsal tubercles are similar sizes in C. candelariensis). Craugastor candelariensis can be differentiated from C. cueyatl and C. hobartsmithi by the absence of vomerine odontophores (present in C. candelariensis). It can be differentiated from C. portilloensis by the presence of posterolateral projections of the frontoparietal (absent in C. candelariensis). Description of holotype. — Holotype small male (SVL ¼ 13.3 mm); snout rounded and short (0.5 mm naris–snout; 4% SVL); long eye–nostril distance (1.7 mm; 13% SVL); tympanum 1.2 mm (7.6% SVL); no supratympanic fold and no shoulder tubercle; finger length formula III ¼ I; single palmar tubercle; single prepollical tubercle; subarticular tubercles present on all fingers; supernumerary tubercles present on Finger III; toe length formula IV Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Variations in paratypes. —Body sizes (SVL) 12.4 mm (MZFC-HE-35617), 14.3 mm (UTA A-66116), 18.6 mm (UTA A-55247); eye–nostril distance 10–13% SVL (males), 9% SVL (female); tympanic ratios 7–10%. Etymology. —The name is an abbreviated allusion to the municipality of Candelaria Loxicha (near the type locality) and the Latin suffix - ensis meaning place. It is simultaneously a reference to the Latin noun candēla meaning a fire or light made of wax, given the translucent yellow appearance of several type specimens in preservative, as if someone were shining a candle through them. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Distribution. —This species is known from intermediate elevations of southern Oaxaca (668–1350 m), an area that mostly consists of Sierra Madre del Sur pine–oak forest habitat. Phylogenetics. — Craugastor candelariensis was strongly supported as monophyletic in the concatenated analysis (ML ¼ 100; BAYES ¼ 1.0; Fig. 3). In this analysis, the sister taxon of C. candelariensis was inferred to be C. polaclavus (ML ¼ 80; BAYES ¼ 0.99). We also observed this sister relationship in the nDNA-only analysis (Fig. 5); however, in the mtDNAonly analysis C. candelariensis was inferred to be the sister taxon of a clade containing C. bitonium þ C. pygmaeus (Fig. 4). In terms of genetic distances, Craugastor candelariensis is most similar to C. polaclavus and C. pygmaeus (both 6.4%; Table 4). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Remarks. —The skull of C. candelariensis is similar to C. bitonium, C. hobartsmithi, C. montanus, and C. pygmaeus, with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. Two type specimens appear white-yellowish in preservative (possibly having been saponified). This species likely co-occurs with C. pygmaeus, C. polaclavus, and C. portilloensis in southcentral Oaxaca (Figs. 6 and 8). In terms of body size and ossification level it is the smallest member of the C. mexicanus series to complete Stage 5 of our ontogenetic sequence.

Craugastor cueyatl sp. nov. Microbatrachylus hobartsmithi: Duellman 1961:33 (in part, based on field IDs EHT-HMS Nos. 18292–18352, includes CAS 87816). [Misidentification]. Eleutherodactylus hobartsmithi: Castro-Franco et al. 2006:107. [Misidentification]. Holotype. — UTA A-62348 (field ID: JAC 27244; Fig. 24B), male collected by J.W. Streicher, C.L. Cox, J. ReyesVelasco, G. Weatherman, and C.M. Sheehy, III, on the road from Avandaro to El Manzano, East of Cerro Gordo, Estado de México, Mexico, 19.12209°N, 100.13969°W, 2311 mon 18 June 2008. Paratypes (2). —MZFC-HE-35614 (Fig. 24A), female with developed ova covered in mildly pigmented connective tissue, same data as holotype, except 19.11735°N, 100.13940°W, 2282 m. AMNH A- 57809 (Fig. 24 C), male from Tepozteco, Morelos, Mexico (Aztec archaeological site, 19.00079°N, 99.10156°W, 2000 m). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Referred specimens (4). — USNM 122054–55, and USNM 139380 from Estado de México and Morelos, respectively. MZFC 1089, from Tepoztlán, Barrio de Ixcaltepec, central Morelos. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 15.7 mm); (2) reduced ossification of the skeleton relative to other members of the series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred, with snout–nostril– canthal–supratympanic stripe; (8) on postrictal tubercle; (9) gular region lightly pigmented; (10) dorsal surface unicolored dark; (11) dark middorsal ridge; (12) evenly tubercular dorsum; (13) body flank unicolored pale with or without a dark supra- or postaxillary band, shagreened; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches anterior corner of eye; (16) outer tarsal ridge with 3–7 rounded tubercles, no raised fringe; (17) finger tips round and not expanded, toe tips slightly lanceolate and barely expanded; (18) similar sizes of inner and outer metatarsal tubercles. Comparisons. — Craugastor cueyatl can be differentiated from C. candelariensis, C. mexicanus, C. montanus, C. omiltemanus, C. portilloensis, and C. saltator by the presence of vomerine odontophores (absent in C. cueyatl). It can be differentiated from C. bitonium, C. hobartsmithi, C. polaclavus, and C. pygmaeus by the absence of posterolateral projections of the frontoparietal (present in C. cueyatl). It can be differentiated from C. rubinus by a larger inner metatarsal tubercle than outer metatarsal tubercle (inner and outer metatarsal tubercles are similar sizes in C. cueyatl). Description of the holotype. — Holotype small male (SVL ¼ 12.3 mm); highly pigmented testes; snout rounded and short (0.6 mm naris–snout; 4% SVL); medium eye– nostril distance (1.4 mm; 11% SVL); tympanum 1.2 mm (10% SVL); supratympanic fold terminating in small shoulder tubercle; finger length formula III Variations in paratypes. —Body sizes (SVL) 15.7 mm (MZFC-HE-35614), 11.7 mm (AMNH A-57809); eye–nostril distance 7% SVL (female), unavailable (male); tympanic ratios 8% (female), 11% (male); crus ratio 51% (female), 54% (male). Etymology. —The specific epithet is taken from the word for frog in Nahuatl, an Aztec language that has been spoken in the Valley of Mexico since the 7th century, a region that includes the type locality of C. cueyatl. Distribution. —The new species occurs throughout the central region of the Trans Volcanic Mexican Cordilleras slopes, those facing the Río Balsas, in the states of Mexico and Morelos (1670–2264 m), an area of mesic pine–oak forest habitat. Castro-Franco et al. (2006) associated C. cueyatl (as E. hobartsmithi) with dry tropical forests and cultivated areas. Diet. —The holotype’s stomach was found to contain a small spider of unknown taxonomy. Phylogenetics. — Craugastor cueyatl was inferred to be the sister taxon of C. bitonium þ C. pygmaeus with strong support in the concatenated analysis (87 ML; 0.99 BAYES; Fig. 3). The placement of C. cueyatl was less certain in the mtDNA and nDNA-only analyses (Figs. 4 and 5). In terms of genetic distances, Craugastor cueyatl is most similar to C. pygmaeus (5.9%; Table 4). Remarks. —The skull of C. cueyatl is similar to that of C. hobartsmithi, C. montanus, and C. pygmaeus with a more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. Several of the referred specimens are based on geographic occurrence and general gestalt and should be further examined. One referred specimen of C. cueyatl (USNM 139380) is reported to be collected from Distrito Federal (¼ Mexico City). We georeferenced this locality as being in the city (Fig. 8), but it is most likely the specimen was collected from outside the metropolis. Very rarely do species of vertebrate occurring near or in major metropolitan areas remain hidden from science; the late discovery of this new species is likely explained by the diminutive size of C. cueyatl and previous confusion with C. hobartsmithi and C. pygmaeus., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 23-25, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Duellman, W. E. 1961. The amphibians of Michoacan, Mexico. Miscellaneous Publications of the University of Kansas Museum of Natural History 15: 1 - 148.","Castro-Franco, R., G. G. Vergara Garcia, M. Guadalupe, B. Zagal, and W. Mena Arizmendi. 2006. Diversidad y distribucion de anfibios del estado de Morelos, Mexico. Acta Zoologica Mexicana 22: 103 - 117. [In Spanish.]"]}

Craugastor omiltemanus (Günther 1900a) Syrrhaphus omiltemanus Günther 1900a:213. Holotype unsexed (BMNH 1901.12.19.7) from ‘‘Omilteme, Guerrero, Mexico.’’ [Examined]. Hylodes calcitrans Günther 1900b:229. Holotype unsexed (BMNH 1901.12.19.25) from ‘‘Omilteme, Guerrero, Mexico.’’ [Examined]. Eleutherodactylus omiltemanus (Günther): Lynch 1970:175. Craugastor omiltemanus (Günther): Crawford and Smith 2005:536. Diagnosis. —Based on Hylodes calcitrans type series of 25 individuals and 1 additional specimen. Aspecies of Craugastor distinguished by the following combination of characters: (1) large adult size (maximum SVL ¼ 38.8 mm); (2) reduced ossification of skeleton in adults relative to other members of series, lacking ossification of any elements beyond Stage 2 (Table 3) except for the sphenethmoid; (3) presence of posterolateral projection of frontoparietal; (4) presence of vomerine odontophore; (5) presence or absence of raised tubercles on eyelids, smooth, six round and only slightly protruding tubercles, sometimes a few aligned at the outer edge; (6) supratympanic fold developed; (7) face flank barred or dark; canthus dark, pale, or spotted; canthus with or without a stripe (complete or broken); (8) one or two postrictal tubercles; (9) gular region from evenly scattered fine pigmentation, to densely pigmented with a mid-pale stripe; (10) dorsal surface unicolored pale brown or grey; unicolored to lightly spotted above the scapular and/or rump areas, or tubercles and ridges (if present), interorbital bar; (11) with or without a middorsal ridge; (12) dorsum smooth with only few fine tubercles toward flanks and urostyle or with ridges forming hourglass patterns and medium tuberculation; (13) body flank darker anteriorly (postaxillary) due to posterolateral expansion of supratympanic stripe; sometimes with contrasting white and dark blotching inguinal area, otherwise pale colored; shagreened; (14) inguinal gland present and axillary gland present in adults; (15) when leg adpressed to body, heel reaches middle of eye to mid-canthal area; (16) outer tarsal ridge 0–5 rounded and only slightly raised tubercles, no raised fringe; (17) finger and toe pads round, fingertips slightly or not expanded, toe tips expanded; (18) inner metatarsal tubercle larger than outer metatarsaltubercle. Comparisons. — Craugastor omiltemanus can be differentiated from all other members of the C. mexicanus series by the combination of rough (and often raised) areolate skin on its venter and a massive inner metatarsal tubercle (Figs. 21G and 29; Lynch 1970, 2000). Description. —In previous literature described as largebodied, short-legged, with Toe III C. omiltemanus, BMNH 1901.12.19.7 (reregistered as BMNH 1947.2.16.62; Fig. 1 J) and BMNH 1901.12.19.8 (reregistered as BMNH 1947.2.16.63). We designate the former as the lectotype and the latter as the paralectotype of this species. Both specimens appear to have partially desiccated at some point in their history, whereas the type series of Hylodes calcitrans is in much better condition (Fig. 1K; Fig. 23A and B). We designate BMNH 1901.12.19.29 (reregistered as BMNH 1947.2.16.47) as the lectotype of H. calcitrans. Relative finger lengthsof the typesare III> IV> II> Iandrelative toe lengths are IV Distribution. —Prior to our study, C. omiltemanus was known only from Guerrero (Günther 1900a; Crawford and Smith 2005). Our discovery of a specimen from Oaxaca (UTA A-64264) that is both phylogenetically and morphologically assignable with C. omiltemanus extends the distribution of this species eastward. This species occurs in high-elevation pine–oak forest habitats of the Sierra Madre del Sur in the states of Guerrero and Oaxaca (Fig. 8; Fig. 30E and F). Phylogenetics. —In all analyses, C. omiltemanus was found to be monophyletic with strong support (ML> 90; BAYES> 0.90; Figs. 3 and 4). In the concatenated analysis C. omiltemanus was found to be the sister taxon of C. saltator, although with limited support (ML ¼ 44; BAYES ¼ 0.74; Fig. 3). There was less support for the sister relationship with C. saltator in the mtDNA-only analysis (ML ¼ 33; BAYES ¼ 0.82; Fig. 4). In the nDNA-only analysis (which did not include C. saltator), C. omiltemanus was placed in basal polytomy with two other clades: (1) C. mexicanus and (2) all other species of the C. mexicanus series (Fig. 5). In terms of genetic distances (Table 4), C. omiltemanus was most similar to C. mexicanus (4.9%), followed by similarity to C. saltator (5.6%). Remarks. —The skull of C. omiltemanus is similar to C. mexicanus and C. saltator with a more anteriorly placed anterior suture of frontoparietal and prootic than that in other species. Skull examination also was conducted on a smaller subadult individual (UTA A-55240, SVL ¼ 16.5 mm) that had only Stage 1 of the ontogenetic sequence complete and only two features of Stage 2 present. Unlike the adult specimen, the nasals are completely unossified and the frontoparietal greatly reduced, and while vomerine odontophores are present the posterolateral projection of the frontoparietal is absent. Craugastor omiltemanus likely shares a similar distribution with other Craugastor species endemic to the Sierra Madre del Sur (e.g., C. uno; Streicher et al. 2011). Lynch (2000) reports that C. omiltemanus has white testes, but we observed pigmented testes in this species (Table 5). Throughout its range, C. omiltemanus likely co-occurs with C. bitonium, C. mexicanus, C. pygmaeus, and C. saltator. Male C. omiltemanus have significantly larger tympana than do female C. omiltemanus (Lynch 2000; this study)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 30-31, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Gunther, A. C. L. G. 1900 a. Reptilia and Batrachia, part 155. Pp. 213 - 220 in Biologia Centrali Americana 7 (O. Salvin and F. D. Godman, eds.). R. H. Porter and Dulau & Co., UK.","Gunther, A. C. L. G. 1900 b. Reptilia and Batrachia, part 157. Pp. 229 - 236 in Biologia Centrali Americana 7 (O. Salvin and F. D. Godman, eds.). R. H. Porter and Dulau & Co., UK.","Lynch, J. D. 1970. Taxonomic notes on some Mexican frogs (Eleutherodactylus: Leptodactylidae). Herpetologica 26: 172 - 180.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94.","Streicher, J. W., J. M. Meik, E. N. Smith, and J. A. Campbell. 2011. Low levels of genetic diversity among morphologically distinct populations of an enigmatic montane frog from Mexico (Craugastor uno: Craugastoridae). Amphibia-Reptilia 32: 125 - 131."]}

Craugastor polaclavus sp. nov. Holotype. — UTA A-62392 (field ID: JAC 21230; Fig. 31), female collected by E.N. Smith and colleagues in Portillo del Rayo, Distrito San Pedro Pochutla, Sierra Madre del Sur, Oaxaca, Mexico, 15.973038N, 96.997118W, 1550–1585 m, 24 September 2001. Paratypes (4). — UTA A- 55246, a recent hatchling collected by E. N. Smith and José Luís Camarillo Rangel from Río Salado, Sierra Madre del Sur, Oaxaca, Mexico, 16.1941678N, 97.09758W, 1245 m, 26 September 1997. UTA A- 66098 and MZFC-HE-35582–83, adult or subadult specimens all collected with the holotype. Referred specimen (1). — UTA A- 66097, female, same data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 14.7 mm); (2) highly reduced ossification of skeleton in adults relative to other members of series; (3) presence of posterolateral projection of the frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred with no distinctive canthal stripe, 1–2 particularly dark bars below eye; (8) one or two postrictal tubercles; (9) gular region with pale spotting; (10) dorsal surface blotched; suprascapular chevron, interorbital bar; (11) pale or as background middorsal ridge; (12) dorsum smooth with only large scattered tubercles; (13) body flank darker anteriorly, no sharp delineation of color change, smooth to shagreened; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches nostril; (16) outer tarsal ridge with 0–4 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not expanded, toe tips slightly expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor polaclavus can be differentiated from C. candelariensis, C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by the presence of vomerine odontophores (absent in C. polaclavus). It can be differentiated from C. bitonium, and C. cueyatl by the condition of adpressed leg where the heel does not reach the nostril (reaches nostril in C. polaclavus). It can be differentiated from C. hobartsmithi and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. polaclavus). It can be differentiated from C. portilloensis by metatarsal tubercles of equal size (unequal in C. polaclavus). It can be differentiated from C. rubinus by relative finger lengths of IV ¼ II (IV> II in C. polaclavus). Description of holotype. — Holotype small female (SVL ¼ 14.7 mm); snout rounded and short (0.8 mm naris–snout; 5% SVL); short eye–nostril distance (1.4 mm; 7% SVL); tympanum 1.4 mm (10% SVL); mild supratympanic fold terminating in small shoulder tubercle; finger length formula III ¼ II Variations in paratypes. —Body sizes in SVL 8.2 mm (UTA A-55246), 12.3 mm (UTA A-66098), 11.6 mm (MZFC-HE-35582), 13.5 mm (MZFC-HE-35583); eye–nostril distance 11–13% SVL; tympanic ratios 7–8%. Distribution. —Intermediate elevations in the foothills of the Sierra Madre del Sur in Oaxaca 1245–1585 m (Fig. 7). These habitats consist of mixed tropical dry and temperate sierra forests. Etymology. —The specific epithet is a combination of the Latin pola meaning small and clavus meaning wart. The name is an allusion to the small size and rugose appearance of several individuals in the type series. Phylogenetics. — Craugastor polaclavus was inferred to be the sister taxon of C. candelariensis with strong support in the concatenated analysis (90 ML; 0.99 BAYES; Fig. 3) and nDNA-only analysis (ML> 90, BAYES> 0.90; Fig. 5). The placement of C. polaclavus was less certain in the mtDNA, where it was found to be the sister taxon of a clade containing C. cf. hobartsmithi þ C. rubinus (ML ¼ 51, BAYES ¼ 0.74; Fig. 4). In terms of genetic distances (Table 4), C. polaclavus was most similar to C. portilloensis (5.8%), followed by similarity to C. bitonium (5.9%). Remarks. —The skull of C. polaclavus is similar to C. mexicanus, C. omiltemanus, and C. saltator, with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. Specimens were dissected; three (UTA A-62392, 66098, and MZFC-HE-35582) seem to be subadult females with unpigmented ovaries and thin undeveloped oviducts, UTA A-66097 is an adult female with a thickened oviduct and also unpigmented ovaries containing yolked eggs, UTA A-66098 adult and MZFC-HE-35582 subadult are males with pigmented testes (smaller on second specimen), and the hatchling (8.2 mm SVL) was not dissected. This species likely co-occurs with C. candelariensis, C. portilloensis, and C. pygmaeus in southcentral Oaxaca (Figs. 6 and 8). It was collected in sympatry with C. portilloensis at Portillo del Rayo, Oaxaca, Mexico (Fig. 8). The hatchling paratype specimen (UTA A-55246) had no evidence of skeletal ossification (Fig. 11)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 32-33, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastor saltator (Taylor 1941) Eleutherodactylus saltator Taylor 1941:89. Holotype female (FMNH 100116) from ‘‘Omilteme, Guerrero, Mexico.’’ [Examined]. Eleutherodactylus mexicanus: Lynch 2000:134. [Misidentification]. Craugastor saltator (Taylor): Crawford and Smith 2005:536. Diagnosis. —Based on holotype and three additional specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) large adult size (maximum SVL ¼ 44 mm); (2) full ossification of skeleton in adults but under a different ontogenetic sequence than other members of the series (Table 3), where the sphenethmoid, humeral interior epiphyses, and tibiofibular epiphyses do not ossify during stage 3, the fontopareital-prootic suture does not offset posteriorly during stage 4, and the epicorocoids do not ossify during stage 6; (3) presence of posterolateral projection of the frontoparietal; (4) presence of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold developed; (7) face flank, labium barred or dark with a cream stripe above; canthal stripe complete or broken; (8) one or two postrictal tubercles; (9) gular region with trace of mid-pale stripe; (10) dorsal surface unicolored, blotched, or with wide middorsal stripe bordered by cream-colored stripes, dark interorbital bar, sometimes with small suprascapular and/or rump spots; (11) middorsal ridge present; (12) dorsum smooth or slightly tuberculate; (13) body flank unicolored, rarely supratympanic stripe extending to area behind insertion of arm, making anterior area darker; finely shagreened; (14) inguinal gland present and axillary gland present in adults; (15) when leg adpressed to body, heel reaches beyond snout; (16) outer tarsal ridge with 0–5 extremely small, flat, and round tubercles, no raised fringe or ridge; (17) finger and toe pads round and expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor saltator can be differentiated from C. bitonium, C. cueyatl, C. hobartsmithi, C. pygmaeus, and C. rubinus by the absence of vomerine odontophores (present in C. saltator). It can be differentiated from C. candelariensis and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. saltator). It can be differentiated from C. omiltemanus by ventral skin texture in life; smooth to granular in C. saltator versus areolate in C. omiltemanus. It can be differentiated from C. montanus and C. polaclavus by shorter relative leg sizes with a crus ratio of 50–58% SVL (long relative leg sizes of 62–73% SVL in C. saltator). Craugastor saltator is most similar to C. mexicanus; however, they do not overlap in geographic range (see C. mexicanus account for additional information). Description. —Detailed description in Taylor (1942). Described as large-bodied, long-legged, with pigmented testes, unequal inner and outer metatarsal tubercle sizes, large vomerine odontophores, and generally smooth dorsal skin (Taylor 1941); with less population-level chromatic variation than its relative C. mexicanus (Lynch 2000). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Holotype (FMNH 100166) large female (SVL ¼ 44 mm; Fig. 36) with relative finger lengths III> IV> II> Iand relative toe lengths IV> III> V> II> I. Asubadult individual used in the molecular analysis, UTA A- 66120 (Fig. 36) had the following characteristics: SVL ¼ 14.5 mm; tympanum width ¼ 1.2 mm; naris–snout length ¼ 0.7 mm (4.7% SVL); eye–nostril distance ¼ 1.5 mm (10.3% SVL); relative finger lengths III> IV> II> I; relative toe lengths IV> III> V> II> I; unlike adult specimens, inner metatarsal tubercle and outer metatarsal tubercle near equal in length. Distribution. —Known only from the high-elevation pine–oak forests of Guerrero in the Sierra Madre del Sur (Fig. 6). Phylogenetics. —We were only able to sequence mtDNA data for C. saltator, so only it was included in the concatenated and mtDNA analyses. In the concatenated analysis, it was recovered as the sister taxon of C. omiltemanus with limited support (ML ¼ 44; BAYES ¼ 0.74; Fig. 3). In the mtDNA-only analysis, support for this relationship was lower in the ML analysis but higher in the BAYES analysis (ML ¼ 33, BAYES ¼ 0.82; Fig. 4). In terms of genetic distances (Table 4), C. saltator was most similar to C. mexicanus (5.1%), followed by similarity to C. omiltemanus (5.6%). Remarks. —The skull of C. saltator is similar to that of C. mexicanus and C. omiltemanus, with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. Taylor (1941:91) makes multiple references to C. saltator and C. omiltemanus (as Eleutherodactylus calcitrans) being similar and states that these two species can be differentiated by the ‘‘very long limb, and the reduced inner metatarsaltubercle [of C. saltator].’’ Lynch (2000) noted that C. saltator is actually far more similar to C. mexicanus from adjacent Oaxaca and subsequently synonymized C. saltator with C. mexicanus (see Methods, Taxonomic History). Despite the finding of Crawford and Smith (2005), which were used to revalidate C. saltator, at one point in our study one of us (JWS) was convinced by Lynch’s (2000) argument that C. saltator should be a junior synonym of C. mexicanus. The basis for this suspicion was (1) a specimen collected by J.D. Godman from Omilteme, Guerrero (the type locality) identified by J.D. Lynch as C. saltator on 13 January 1972 (BMNH 1901.12.19.24) is similar in morphology to C. mexicanus; (2) the nDNA analysis of Crawford and Smith (2005) found C. mexicanus þ C. saltator to be monophyletic; and, (3) we did not observe range overlap between C. mexicanus and C. saltator (Fig. 6), which may be consistent with a single large-bodied, long-legged species inhabiting the Sierra Madre del Sur. However, as in Crawford and Smith (2005), our mtDNA phylogenetic results did not recover C. mexicanus þ C. saltator as monophyletic (Fig. 3) and the specimens of C. saltator we examined had on average larger body sizes and differing toe length formulae than did C. mexicanus (Figs. 10 and 11; Table 6). As such, we continue to recognize C. saltator as a distinct species pending further taxonomic investigation. Craugastor saltator likely co-occurs with C. bitonium, C. pygmaeus, and C. omiltemanus in the Sierra Madre del Sur of Guerrero. It may overlap with C. mexicanus in eastern Guerrero and western Oaxaca (Fig. 6)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 37-38, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Taylor, E. H. 1942. New tailless Amphibia from Mexico. University of Kansas Science Bulletin 28: 67 - 89."]}

Craugastor portilloensis sp. nov. Holotype. — UTA A-62393 (field ID: JAC 21431; Fig. 32), subadult female collected by E.N. Smith and colleagues in Portillo del Rayo, Distrito San Pedro Pochutla, Sierra Madre del Sur, Oaxaca, Mexico, 15.9794448N, 96.5166678W, 1550 m, 1 October 2001. Paratypes (4). —MZFC-HE-35580 and UTA A- 66095, juvenile males. MZFC-HE- 35581 subadult female, and UTA A- 66096 subadult male. All collected at the type locality between 1550 and 1585 mon 24 September 2001 by E.N. Smith and colleagues. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 11.4 mm); (2) reduced ossification of skeleton in adults relative to other members of the series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank darker than dorsum, faintly barred lips, snout–nostril– canthal–supratympanic stripe; (8) one or two postrictal tubercles; (9) gular region with pale spotting; (10) dorsal surface unicolored pale, diffuse interorbital bar, sometimes with two dark rump spots; (11) with or without a middorsal ridge; (12) dorsum smooth or with scattered fine tubercles; (13) body flank has dark supratympanic stripe extending toward lower mid-flank, smooth, very few small tubercles; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and slightly past snout; (16) outer tarsal ridge smooth or with 1–3 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not expanded, toe tips slightly expanded; (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor portilloensis can be differentiated from C. candelariensis, C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by the presence of vomerine odontophores (absent in C. portilloensis). It can be differentiated from C. bitonium, C. hobartsmithi, and C. pygameus by the absence of a posterolateral projection of the frontoparietal (present in C. portilloensis). It can be further differentiated from C. pygmaeus by the absence of a canthal mask (present in C. portilloensis). It can be differentiated from C. cueyatl and C. polaclavus by metatarsal tubercles of unequal size (equal in C. portilloensis). It can be differentiated from C. rubinus by relative finger lengths of IV ¼ II (IV> II in C. portilloensis). Description of holotype. — Holotype small female (SVL ¼ 11.4 mm); snout rounded and short (0.8 mm naris–snout; 6% SVL); short eye–nostril distance (1.1 mm; 9.8% SVL); tympanum 0.8 mm (6.8% SVL); no supratympanic fold; small shoulder tubercle; finger length formula III Variations in paratypes. —Body sizes in SVL 8.7 mm (MZFC-HE-35580), 8.4 mm (UTA A-66095), 12.1 mm (MZFC-HE-35581), 12.6 mm (UTA A-66096); eye–nostril distance 10–11% SVL; tympanic ratios 7%. Etymology. —The name is an abbreviated allusion to the type locality near the town of Portillo del Rayo and the Latin suffix - ensis meaning place. Distribution. —Intermediate elevations in the foothills of the Sierra Madre del Sur in Oaxaca 1550–1585 m (Fig. 8). These habitats consist of mixed tropical dry and temperate sierra forests. Phylogenetics. —In the concatenated analysis, Craugastor portilloensis is the sister taxon to a clade of small-bodied species (C. bitonium, C. candelariensis, C. cueyatl, C. hobartsmithi, C. polaclavus, C. pygmaeus, and C. rubinus), but with low branch support (ML ¼ 53, BAYES ¼ 66; Fig. 3). It occupies the same phylogenetic placement in the mtDNA-only data set (Fig. 4), but in the nDNA-only data set is the sister taxon of a clade including (C. bitonium, C. cueyatl, C. hobartsmithi, C. pygmaeus, and C. rubinus) with moderate support in the Bayesian analysis (ML ¼ 43, BAYES ¼ 0.78; Fig. 5). In terms of genetic distances (Table 4), C. portilloensis was most similar to C. montanus (5.7%), followed by similarity to C. polaclavus (5.8%). Remarks. —The skull of C. portilloensis is similar to C. mexicanus, C. omiltemanus, and C. saltator, with a more anteriorly placed anterior suture of frontoparietal and prootic than in other species. We dissected all specimens of C. portilloensis; males possess pigmented testes and females unpigmented ovaries. This species likely co-occurs with C. candelariensis, C. polaclavus, and C. pygmaeus in southcentral Oaxaca (Figs. 6 and 8). It was collected in sympatry with C. polaclavus at the type locality of Portillo del Rayo, Oaxaca, Mexico (Fig. 8)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 33-34, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastormexicanus (Brocchi 1877) Leiuperus mexicanus Brocchi 1877:184. Holotype unsexed (MNHNP 6218) from ‘‘ Mexico ’’ (¼ Cerro San Felipe, Oaxaca, Mexico [Smith and Taylor 1950]). [Examined]. Paludicola mexicana (Brocchi): Boulenger 1882:237; Neiden 1923:513. Pleurodema mexicana (Brocchi): Parker 1927:475. Microbatrachylus oaxacae Taylor 1940:505. Holotype male (FMNH 100001) from ‘‘Cerro San Flipe, near Oaxaca, Oaxaca, Mexico.’’ [Examined]. Microbatrachylus lineatissimus Taylor 1941:87. Holotype male (FMNH 1000036) from ‘‘Cerro San Flipe, near Oaxaca, Oaxaca, Mexico.’’ [Examined]. Microbatrachylus fuscatus Davis and Dixon 1957:146. Holotype female (TCWC 12171) from ‘‘ 20 miles east of Tulancingo, Hidalgo, Mexico.’’ [Not examined; neartopotypic specimen from Hidalgo examined (UTA A-66138)]. Eleutherodactylus oaxacae (Taylor): Lynch 1965:3. Eleutherodactylus lineatissimus (Taylor): Lynch 1965:3. Eleutherodactylus mexicanus (Brocchi): Gorham 1966:86. Craugastor mexicanus (Brocchi): Crawford and Smith 2005:536. Diagnosis. —Based on 26 specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) large adult size (maximum SVL ¼ 40.5 mm); (2) full ossification of the skeleton in adults; (3) presence of posterolateral projection of frontoparietal (Fig. 26B); (4) presence of vomerine odontophores (in larger individuals); (5) presence or absence of raised tubercles on eyelids, Comparisons. — Craugastor mexicanus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by the equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. mexicanus). It can be differentiated from C. bitonium and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. mexicanus). It can be differentiated from C. polaclavus and C. rubinus by the absence of vomerine odontophores (present in C. mexicanus). It can be differentiated from C. montanus by the condition of supratympanic folds in adults; developed in C. mexicanus versus moderate to poorly developed in C. montanus. It can be differentiated from C. omiltemanus by ventral skin texture in life; smooth to granular in C. mexicanus versus areolate in C. omiltemanus. Craugastor mexicanus is most similar to C. saltator. We were unable to identify any reliable morphological characters to differentiate C. mexicanus and C. saltator; however, they have nonoverlapping geographic distributions, with C. saltator only occurring in western Guerrero (Fig. 6). Description. —In previous literature, described as largebodied, long-legged with row of small tubercles on outer edge of the tarsus (Taylor 1941); variable palmar tubercle arrangements (palmar tubercle divided, single, or evaginated; Lynch 1965); inner metatarsal tubercle larger than outer metatarsal tubercle; lack of tarsal tubercles (Lynch 2000). Holotype (MNHNP 6318) is large (~ 40 mm SVL); owing to poor preservation, columella of holotype partially ruptured the tympana on both sides (Fig. 1B); finger lengths III> IV> II> I; toe length IV> V ¼ III> II> I. Distribution. —This species is widespread throughout eastern Mexico in high-elevation habitats (1554–2700 m) of Oaxaca, Puebla, Hidalgo, and Veracruz (Fig. 6). These habitats span the Sierra Madre Oriental and Sierra Madre del Sur. Canseco Márquez and Gutiérrez Mayén (2010) report that C. mexicanus occurs in the forests adjacent to the Tehuacán-Cuicatlán Valley in Puebla and Oaxaca. It is likely this species occurs in Guerrero; however, most specimens resembling C. mexicanus we examined from Guerrero are referrable to C. saltator. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Phylogenetics. —The concatenated data set placed C. mexicanus as the sister taxon to a clade of C. omiltemanus þ C. saltator with high support in the BAYES analysis (0.98) but moderate support in the ML analysis (75; Fig. 3). This appears to be a relationship supported exclusively by the mtDNA data set because separate analysis of the nDNA markers did not confidently infer a sister taxon of C. mexicanus (Figs. 4 and 5). Although BAYES analyses strongly supported the monophyly of C. mexicanus (> 0.90), the ML analyses recovered variable support ranging from 79 (concatenated analysis) to 54 (nDNA-only analysis). In terms of genetic distances (Table 4), C. mexicanus was most similar to C. omiltemanus (4.9%), followed by similarity to C. saltator (5.1%). Intraspecific variation. —We examined over 220 specimens of C. mexicanus for this revision (Appendix I), and briefly provide some patterns of intraspecific variation that we observed. Many populations have substantial colorpattern polymorphism (Fig. 27), similar to what is observed in the C. rhodopis series (Lynch 1966; Streicher et al. 2014). In some northern populations (Hidalgo and Puebla), the canthal mask is broken into a black spot posterior to the tympanum (often with a thin yet distinctive white margin). Specimens throughout Oaxaca varied in whether they had a single or divided palmar tubercle (as reported by Lynch 1965, see his Fig. 2); in six specimens we examined there was asymmetry with a single palmar tubercle on one hand and divided on the other. Patterns of dorsal ridging and coloration often coincide, with observations of the following morphs: (1) straight raised ridges that are tubercular and darker in coloration than the rest of the dorsum; (2) straight raised dorsal ridges that are tubercular but of the same color as the rest of the dorsum; (3) straight lines of color that are dark but with no tubercular raised ridges; (4) ridges that are not straight but form an hourglass shape and are tubercular; (5) ridges that are not straight but form an hourglass shape and are not tubercular; (6) scattered tubercles on the dorsum, which are sometimes dark and sometimes same color as the background; (7) pale or dark unicolored dorsum; (8) a unicolored dorsum with a pale median stripe that can be ridged or smooth; (9) a wide dark band on the dorsum, which usually co-occurs with a pale upper labium; and, (10) white hills of coloration and a peppered grayish dorsum sometimes with a diffused lighter cream color, a pattern that may mimic bird droppings (appearing superficially as a smear of uric acid and digestive waste). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Remarks. —The skull of C. mexicanus is similar to that of C. omiltemanus and C. saltator, with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. We noticed that the ‘‘ M. lineatissimus ’’ morphotype (raised and parallel ridges on the dorsum) occurs widely throughout the range of C. mexicanus. Interestingly, the examined paratype of ‘‘ M. lineatissimus ’’ has a unique skull shape for C. mexicanus. The shape is similar to that of C. hobartsmithi, C. montanus, and C. pygmaeus, with a more posteriorly placed anterior suture of the frontoparietal and prootic than in other species, coupled with a narrower back of the skull than in any other species (Fig. 27A). This unique condition likely explains the specimen as an outlier in several of our statistical analyses (Figs. 9, 11, and 12). However, we were unable to CT-scan other individuals with the ‘‘ M. lineatissimus ’’ morphotype (or the holotype of M. lineatissmus), so future investigation is necessary to determine whether the examined paratype (FMNH 104548) is an aberrant individual of C. mexicanus or represents a valid taxon. Lynch (1965) reports that ‘‘ E. oaxacae ’’ has parotoid glands. However, we saw no evidence of these glands in the two type specimens that we examined (FMNH 100001 and FMNH 126638), nor are we aware of any species of Craugastor that possess parotoid glands (¼ large poison glands on the nuchal region). Craugastor mexicanus likely co-occurs with C. omiltemanus at high-elevation localities of central Oaxaca (Figs. 6 and 8). At intermediate elevation localities in Veracruz and Oaxaca, it may overlap with C. pygmaeus (Fig. 7). Male C. mexicanus have significantly larger tympana than do female C. mexicanus (Fig. 13)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 27-29, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Brocchi, P. 1877. Sur quelques batraciens raniformes et bufoniformes de l'Amerique Centrale. Bulletin de La Societe Philomathique de Paris 1: 175 - 179. [In French.]","Smith, H. M., and E. H. Taylor. 1950. Type localities of Mexican reptiles and amphibians. The University of Kansas Science Bulletin 33: 313 - 339.","Boulenger, G. A. 1882. Catalogue of the Batrachia, Salientia and Ecaudata in the Collection of the British Museum. Natural History Museum Publications, UK.","Neiden, F. 1923. Anura. I. Subordo Aglossa und Phaneroglossa, Sectio 1. Arcifera. Das Tierreich 46: 32 - 584. [In German.]","Parker, H. W. 1927. A revision of the frogs of the genera Pseudopaludicola, Physalaemus, and Pleurodema. Journal of Natural History 20: 450 - 478.","Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571.","Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94.","Davis, W. B., and J. R. Dixon. 1957. Notes on Mexican amphibians, with description of a new Microbatrachylus. Herpetologica 13: 145 - 147.","Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Canseco Marquez, L., and M. G. Gutierrez Mayen. 2010. Anfibios y Reptiles del Valle Tehuacan-Cuicatlan. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Mexico.","Lynch, J. D. 1966. Multiple morphotypy and parallel polymorphism in some neotropical frogs. Systematic Biology 15: 18 - 23.","Streicher, J. W., U. O. Garcia-Vazquez, P. Ponce-Campos, O. Flores-Villela, J. A. Campbell, and E. N. Smith. 2014. Evolutionary relationships amongst polymorphic direct-developing frogs in the Craugastor rhodopis Species Group (Anura: Craugastoridae). Systematics and Biodiversity 12: 1 - 22."]}

Craugastor bitonium sp. nov. Holotype. — UTA A-64254 (field ID: JAC 22117; Fig. 22A), adult female from road between Yerba Santa and Yextla (HWY 196), Guerrero, Mexico, 17.52666°N, 99.9579°W, 2071 m, collected by J.A. Campbell and colleagues on 10 June 2002. Paratypes (5). —MZFC-HE-35600–01 and UTA A- 66117– 18 adult females, and UTA A-66119 adult male (Fig. 22B–D), all same collection data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 16.7 mm); (2) full ossification of skeletal elements in adults; (3) absence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred or with supralabial pale stripe, and with or without dark canthal stripe; (8) single postrictal tubercle; (9) gular region peppered with melanocytes; (10) dorsal surface two-toned, usually with a dark suprascapular ^ shape, or almost unicolored; (11) pale or ground color middorsal ridge; (12) scattered fine tubercles on dorsum; (13) body flank barred darker anteriorly, slightly shagreened to smooth; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches middle of eye to slightly beyond snout; (16) outer tarsal ridge with 1–6 small mostly round tubercles, no raised fringe; (17) finger and toe tips round, slightly lanceolated, slightly expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor bitonium can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by their large adult body sizes of SVL> 20 mm (C. bitonium), the presence of vomerine odontophores (absent in C. bitonium), and the presence of three palmar tubercles (one palmar tubercle in C. bitonium). Craugastor bitonium can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by the presence of metatarsal tubercles of similar sizes (different sizes in C. bitonium). Craugastor bitonium can be differentiated from C. rubinus by the presence of posterolateral projections of the frontoparietal (absent in C. bitonium). Craugastor bitonium can be differentiated from C. polaclavus by its shorter eye–nostril distance with an eye–nostril distance 9–10% SVL in C. bitonium and 10–13% SVL in C. polaclavus. Craugastor bitonium is most similar to C. pygmaeus (in morphology, osteology, and genetic distance), but may be differentiated from this taxon by the condition of the outer tarsal ridge; 1–6 small tubercles in C. bitonium versus no tubercles (¼ smooth) in C. pygmaeus. Holotype description. — Holotype small female with unpigmented developing ova (SVL ¼ 15.8 mm); snout rounded and short (0.9 mm naris–snout; 6% SVL); short eye–nostril distance (1.42 mm; 9% SVL); tympanum 1.4 mm (8.9% SVL); small supratympanic fold terminating in shoulder tubercle; finger length formula III Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Variations in paratypes. —Body sizes (SVL) 15.8 mm (MZFC-HE-35600), 15.2 mm (MZFC-HE-35601), 16.9 mm (UTA A- 66117), 16.7 mm (UTA A- 66118), 12.3 mm (UTA A- 66119); eye–nostril distance 9–10% SVL; tympanic ratios 7– 9%; dorsal color patterns variable, often with two distinctive patches of differing ground coloration ranging from orange to tan. Etymology. —The specific epithet is a combination of the Latin prefix bi- meaning two and tonium meaning tone. It is a reference to the two distinctive patches of color found on the holotype and several paratypes that create the appearance of a ‘‘two-tone’’ dorsal coloration. Distribution. —This species is known only from the Sierra Madre del Sur of central Guerrero (~ 2071 m). The closest named places to the type locality are Izotepec to the north and Los Bajos to the southwest. The habitat at the type locality is montane pine–oak forest. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Diet. —CT-scan of the holotype revealed the presence of a small millipede (Diplopoda) in the stomach. We also noted the presence of a small red ant (Formicidae) in the mouth of the holotype. Phylogenetics. — Craugastor bitonium was inferred to be the sister taxon of C. pygmaeus, with high support in the concatenated analyses (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). This sister relationship was also recovered in both mtDNA and nDNA analyses, although with lower support in the nDNAonly analyses (ML ¼ 54, BAYES ¼ 0.67; Figs. 4 and 5). Craugastor bitonium is separated from C. pygmaeus by a P - distance of 4.7% (Table 4). Remarks. —The skull of C. bitonium is similar to C. hobartsmithi, C. montanus, and C. pygmaeus. Craugastor bitonium displays a developmental pattern similar to C. pygmaeus (with high levels of ossification at small sizes), suggesting small adult body sizes (Fig. 11). This species likely co-occurs with C. pygmaeus, C. omiltemanus, and C. saltator in central Guerrero (Figs. 6 and 8). The specimen UTA A-66132 is referred with some hesitation because it was collected from a lower elevation than the type locality and has a Finger Ilength nearing C. pygmaeus (which also occurs in Guerrero). Six of the specimens were adult females containing unpigmented ovaries with yolked eggs and thick oviducts, the seventh (UTA A-66119; Fig. 22C, far left) was an adult male with pigmented testes.

Megophrys acehensis sp. nov. (Figs. 4 A−D, 5A−E) Holotype MZB. Amph 26098 (field number ENS 18664; GenBank accession no. MT 710708; Figs. 4 A−B, 5A−E; Megophrys sp. north in Fig. 1 and Table 1), adult male collected at 2330 h on 5 August 2015 from Aceh Province, Aceh Tengah Regency, Linge District, Kute Robel (4.506444 °N, 96.860750 °E, 1638 m a.s.l), by Elijah Wostl, Ilham Fonna, and Muhammad Iksan (Fig. 6). Paratype UTA A–66178 (field number ENS 21030; GenBank accession no. MT 710709; Fig. 4C–D), a sub adult male collected on at 1856 h on 1 June 2016 from Aceh Province, Pidie Regency, Geumpang District, UPT V Geumpang (4.854540 °N, 96.216540 °E, 1086 m a.s.l) by Michael B. Harvey and Eric N. Smith (Fig. 6). Etymology. The specific name acehensis is derived from the province of Aceh in northern Sumatra and the Latin suffix– ensis meaning from that place. Suggested English common name. Aceh Horned Frog Suggested Indonesian name. Katak–tanduk Aceh Diagnosis. The new species was assigned to the genus Megophrys based on the combination of the following morphological characters, as defined by Kuhl and van Hasselt (1822) and Delorme et al. (2006): (1) pointed snout profile, bearing a pointed projection, protruding laterally beyond the lower jaw; (2) broad and flattened eyelid with palpebral projection; (3) possession of a broad and depressed head; (4) conical spine at the corner of mouth; (5) vertical pupil; (6) presence of maxillary and vomerine teeth. Megophrys acehensis sp. nov. can be diagnosed from its geographically relevant congeners in the Sunda Shelf and the Philippines by the following combination of characters: medium body size, stocky (SVLh 38.1 mm in adult male); snout pointed with a short, acute rostral appendage (RSAL 1.3% in adult male); relatively short triangular palpebral projection with acute tip (EHL 25.1% UEWh in adult male); head length relatively short (RHLh 37.2% in adult male); head wider than its long (HW 113.6% HLh in adult male); tympanum distinct, about one-third of eye diameter (TDH 35.6% ED in adult male), nearly rounded (TDH 85.9 % TDV in adult male); vomerine teeth present; a pair of dorsolateral folds, extending from shoulder, above axilla to groin; dorsal and lateral skin tuberculate, shagreened on the throat to belly; short lower arm (RLAL 45.8% in adult male); foot nearly as long as thigh (FL 96.6% TL in adult male); tibiotarsal articulation reaching posterior of eye; toe webbing absent. Description of holotype (measurements in mm). Adult male, medium body size (SVLh 38.1, SVL 37.4) and habitus stocky; head depressed and broad, wider (HW 16.4, 42.9% SVLh) than long (HL 13.3, 34.9% SVLh); snout short (SL 4.2, 11.1% SVLh), pointed at tip with acute short rostral appendage (SAL 0.7, 1.8% SVLh), laterally protruding and projecting beyond lower jaw; nostril positioned laterally, near to snout than to eye; eye positioned laterally, large, nearly three times of tympanum horizontal diameter (ED 5.5, 280.5% TDH), eye diameter slightly wider (ED 5.5, 14.4% SVLh) than snout–horn length (SLh 5.4, 14.2% SVLh), about two and three-quarter times of nostril–eye length (ED 275% NEL), pupil vertical elliptical; canthus rostralis with sharp, angular ridge, lore sloping and concave; internarial distance (IND 4.0, 10.4% SVLh) about two-thirds of interorbital distance (IND 68.4% IOD); palpebral projection length about one-quarter of total upper eyelid width (EHL 1.2, 25.1% UEWh), tip acute, surface smooth and scattered with small and low tubercles; tympanum distinct, smooth, oval, slightly rounded (TDV 2.3, 6.0% SVLh; TDH 2.0, 5.1% SVLh; TDH 85.9% TDV); angular supratympanic fold, distinct, widened anteriorly, narrowed posteriorly, extending from behind eye, curving down around upper border of tympanum and ending above axilla; white conical tubercles behind the supratympanic fold and anterior to axilla; spinous gland on corner of mouth on jaw angle; single row of maxillary teeth present; vomerine teeth in two widely separated groups, at level posterior borders of choana; tongue lanceolate, notched posteriorly, without papillae; median subgular vocal sac present, having slit–like opening on each side of jaw commissures. * Fold forming Y, X or H on the parietoscapular region to the level of axilla; ** Dorsolateral fold shape: dorsolateral folds are elongated and extend from the parietoscapular region to the groin (Type I); dorsolateral folds extend from the central of parietoscapular region to mid-body (Type II); multiple dorsolateral folds - at least three or four - and they are discontinuous, formed by a series of elongated tubercles (Type III); dorsolateral folds are elongated and curve from the axillary region towards (and reaching) the posterior dorsal margin of tympanum (Type IV). Forelimb slender and short, hand length about half of arm length (HAL 9.5, 54.3% LAL), lower arm proximally enlarged, wider than upper arm; fingers moderately slender, with rounded and swollen tips, unwebbed and lacking of lateral fringes; finger length formula I Hindlimb slender, moderately long (HLL 52.0, 136.5% SVLh), thigh (FML 17.1, 44.8% SVLh) slightly longer than tibia (TL 15.4, 40.4% SVLh), about twice of tarsus length (TSL 8.5, 22.3% SVLh), and nearly as long as foot (FL 14.9, 39.1% SVLh); toe length formula I Dorsal skin surface tuberculate, low dense tubercles in entire dorsal and larger-sized tubercles mostly on forelimb, hindlimb, flanks and groin, shagreened in preservative; a pair of dorsolateral folds present, one on each side, extending from shoulder above the axilla to groin; vent with dorsal dermal accessory extension; transverse folds on limbs present, three on lower arm, three on thigh and three on tibia; limbs skin laterally tuberculate, larger size of tubercles than on dorsal side; ventral skin smooth shagreened on throat to chest, but smooth on belly and limbs, wrinkled in preservative; a pair of white conical pectoral glands, at base of axilla; white conical femoral glands, at the mid flanks of the posterior thigh; dark brown microspinules nuptial pads covering dorsal and median surfaces of the distal half of metacarpal and proximal half of basal phalanx of first and second fingers. Coloration. In life (Fig. 4A–B, 5A–B), pupil dark, iris golden–brown, base of dorsum immaculate dark brown with indistinct dark brown inverted triangle pattern on parieto-orbital to scapular region and inverted “V” like pattern on back, positioned closer to vent than to parietal region; dorsal surface of head, forelimb and hindlimb darker than lateral body; dark brown transverse folds present on dorsal surface of forearm, thigh and tibia, two on the forearm, three on thigh and three on tibia, fingers with dark brown crossbars, one on first and second fingers, two on third and fourth fingers; dorsomedial surfaces of first and second fingers with dark brown microspinular nuptial pads; knee with irregular dark brown spots; dorsal surface and lateral sides of head dark brown, irregular lighter brown pattern below canthus rostralis, light brown on underside of eyelids; body flanks dark brown and unmarked, lighter than dorsum; forelimb flanks dark brown, as dorsum; hindlimb flanks cream with dark brown blotches; ventral surface from throat to chest dark brown with heavy dark longitudinal marking; belly and underside of fore- and hindlimbs light cream with dark brown blotches; in preservative, pattern remains, but dorsal, lateral, and ventral surfaces are darker (Fig. 5C). Variation. The two type specimens are morphometrically similar. The holotype has slightly longer rostral appendage, head, and snout-nostril and snout lengths, but a slightly shorter palpebral projection, and a thicker and narrower head. Morphometric variation is shown in Table 2. The transverse fold in the occipital region varies in degree of development, being distinct on the holotype but indistinct on the paratype. The dorsal coloration in life is dark brown on the holotype and orange-brown on the paratype. The ventral color of the two specimens is marbled with brown in both specimens but, the background color of the holotype is cream anteriorly and light grey posteriorly while that of the paratype is dark orange anteriorly and whitish posteriorly (Fig. 4 A−D). Comparisons. Megophrys acehensis sp. nov. differs from its most similar congener, M. parallela by having dorsolateral folds extending to the groin (vs. maximum of two-thirds length of trunk: Inger and Iskandar, 2005; Munir et al., 2018), a rostral appendage present (vs. absent: Inger and Iskandar, 2005; Munir et al., 2018), shagreened ventral surface from throat to belly (vs. smooth: Inger and Iskandar, 2005; Munir et al., 2018), and a relatively shorter foot to thigh ratio—FL 92.1−96.6% TL (vs. 105.4−112.5% TL: Munir et al., 2018). From M. montana, the new species differs by having tuberculate dorsal skin surface (vs. smooth: Munir et al., 2018), shagreened skin on throat (vs. smooth: Munir et al., 2018), a relatively short foot to thigh ratio—FL 92.1−96.6% TL (vs. FL 106.2−115.6% TL: Munir et al., 2018), and absent toe webbing (vs. rather developed webbing from second to fifth toes: Munir et al., 2018). From Megophrys selatanensis sp. nov., Megophrys acehensis sp. nov. differs by having a longer rostral appendage—SAL 1.3–1.8% SVLh (vs. SAL 0.3−0.6% SVLh), a slightly longer palpebral projection —EHL 25.1–28.4% UEWh (vs. EHL 21.3–24.3% UEWh), slightly narrower ratio of head width to its length—HW 114.1−115.3% HLh (vs. HW 119.3−126.4% HLh), wider ratio of tympanum to eye diameter—TDH 35.6−37.5% ED (vs. TDH 27.3−28.5% ED), tympanum nearly rounded—TDH 85.9–93.8 TDV (vs. vertically elongated, TDH 46.5−61.9% TDV), slightly shorter lower arms—LAL 44.8–45.8% SVLh (vs. LAL 48.0–49.2% SVLh), tuberculate dorsal skin surface (vs. smooth), shagreened skin on throat (vs. smooth), and absent toe webbing (vs. toes webbed at base). Megophrys acehensis sp. nov. differs from M. lancip by males having a shorter rostral appendage—SAL 1.3−1.8% SVLh (vs. SAL 2.9−3.1% SVLh in males, but overlapped with females SAL 1.4−4.1% SVLh: Munir et al., 2018), shorter head length—HLh 37.2–37.8% SVLh (vs. HLh 42.1–44.6% SVLh; Munir et al., 2018), wider head in males—HW 114.1–115.3% HLh (vs. HW 103.2–105.8% HLh in males, but overlapped with females HW 110.6–116.0% HLh: Munir et al., 2018), wider tympanum relative to eye diameter—TDH 35.6−37.5% ED (vs. TDH 25.7−28.0 ED: Munir et al., 2018), tympanum nearly rounded—TDH 85.9–93.8 TDV (vs. vertically elongated, TDH 61.1−69.1% TDV), relatively short foot to thigh ratio—FL 92.1−96.6% TL (vs. FL 102.5−109.8% TL; Munir et al., 2018), and toe webbing absent (vs. rather developed on first to fifth toes, see Figs. 4D, I, 5B in Munir et al., 2018). Megophrys acehensis sp. nov. differs from M. nasuta by the absence of additional lateral flank folds (vs. presence: Munir et al., 2018, 2019), males being smaller—known adult male 37.4 mm SVL (vs. SVL 66.0– 93.4 mm: Munir et al., 2019), shorter acute rostral appendage—SAL 1.3−1.8% SVLh (vs. acuminate rostral appendage, SAL 1.8−9.2% SVLh: Munir et al., 2019), and shorter acute palpebral projection—EHL 25.1−28.4% UEWh (vs. acuminate palpebral projection, EHL 32.7−61.4% UEWh: Munir et al., 2019). From M. kalimantanensis, the new species differs by the absence of additional flank folds (vs. present: Munir et al., 2019), smaller body size in males—known adult male 37.4 mm SVL (vs. SVL 64.3–100.6 mm: Munir et al., 2019), slightly longer rostral appendage—SAL 1.3−1.8% SVLh (vs. SAL 0.1−0.8% SVLh: Munir et al., 2019), and shorter palpebral projection—EHL 25.1−28.4% UEWh (vs. 30.9−53.2% UEWh: Munir et al., 2019). From M. stejnegeri, the new species differs by having dorsolateral folds extending to the groin (vs. maximum of two-thirds the length of the trunk: Inger 1954), males being smaller—known adult male 37.4 mm SVL (vs. adult male 40.6−60.0 mm: Taylor, 1920; Inger, 1954), vomerine teeth present (vs. absent: Taylor, 1920; Inger et al., 1954), and smooth skin on the posterior of tympanum (vs. tuberculate, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016). Megophrys acehensis sp. nov. can be distinguished from M. kobayashii by having type I dorsolateral folds (vs. type II, see Table 3), absent additional lateral flank folds (vs. present: Malkmus and Matsui, 1997; Munir et al., 2018), males being smaller—known adult male 37.4 mm SVL (vs. 93.0−101.0 mm: Inger and Stuebing, 2005), rostral appendage present (vs. absent: Malkmus and Matsui, 1997), and smaller and fewer tubercles on flanks (vs. larger and numerous: Malkmus and Matsui, 1997). From M. ligayae, the new species differs by having type I dorsolateral folds, extended to the groin (vs. type II, reaching a maximum of two-thirds of trunk: Taylor, 1920; Inger, 1954; Munir et al., 2018), absent additional lateral folds (vs. present: Taylor, 1920; Inger, 1954; Munir et al., 2018), males being smaller—known adult male 37.4 mm SVL (vs. SVL 60.4−69.0 mm: Taylor, 1920; Inger, 1954), and toe webbing absent (vs. rather developed web on third, fourth, and fifth toes: Diesmos et al., 2015). Megophrys acehensis sp. nov. differs from M. edwardinae by having type I dorsolateral folds (vs. absent: Inger 1989), a rostral appendage present (vs. absent: Inger 1989), vomerine teeth present (vs. absent: Inger 1989), and slightly shorter thighs—TL 40.4−41.8% SVLh (vs. TL 45.0−50.0% SVL: Inger 1989). Megophrys acehensis sp. nov. can be distinguished from M. baluensis by having type I dorsolateral folds (vs. type III: Munir et al., 2018), males being slightly smaller—known adult male 37.4 mm SVL (vs. SVL 41.0−45.0 mm: Inger et al., 1995), a rostral appendage present (vs. absent: Boulenger, 1899; Inger, 1966; Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger, 1989; Inger et al., 1995), having smaller and fewer tubercles on body flanks (vs. larger and numerous: Inger et al., 2017), and adpressed tibiotarsal articulation reaching posterior of eye (vs. shoulder: Boulenger, 1899). Megophrys acehensis sp. nov. can be distinguished from M. dringi, M. aceras and M. longipes by the absence of Y, X,>–et al., 1995, see Table 3). Furthermore, from M. dringi, the new species being smaller in male—known adult male 37.4 mm SVL (vs. SVL 43.0−47.0 mm: Inger et al., 1995), having a stocky body (vs. slender: Inger et al., 1995), a rostral appendage present (vs. absent: Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger et al., 1995), vomerine teeth present (vs. absent: Inger et al., 1995), distinct tympanum (vs. partially obscured: Inger et al., 1995). From M. aceras, the new species differs by having smaller body size in male—known adult male 37.4 mm SVL (vs. SVL 48.0− 62.4 mm: Inger and Iskandar, 2005), a stocky body (vs. slender: Boulenger, 1903), rostral appendage present (vs. absent: Boulenger, 1903), tibiotarsal articulation reaching to posterior corner of eye (vs. shoulder, angle of jaws or temporal area: Taylor, 1962), toes web absent (vs. rather developed web on third, fourth, and fifth toes: Taylor, 1962, see figure 5 in Munir et al., 2018). From M. longipes, the new species differs by having a smaller body size in male—known adult male 37.4 mm SVL (vs. SVL 38.9−45.2 mm: Inger and Iskandar, 2005), having a stocky body (vs. slender: Boulenger, 1885), a rostral appendage present (vs. absent: Boulenger, 1885), having a triangular palpebral projection (vs. small like tubercle: Boulenger, 1885; Taylor, 1962), shorter thigh—TL 0.41−0.42 SVL (vs. TL 0.55−0.60 SVL: Inger and Iskandar, 2005) and tibiotarsal articulation reaching posterior corner of eye (vs. far beyond tip of snout; Boulenger 1885). Distribution and Natural History. The holotype of Megophrys acehensis sp. nov. was collected on leaf litter in a sloping area at the edge of a primary forest near a stream, while the paratype was collected from leaf litter in a palm oil plantation near the edge of an old secondary forest. Landslides, new road development, and monoculture forests have become major threats at the holotype locality, while the threats at the paratype locality were the land use changes and water pollution from palm oil fields. The precise distribution, population, habitat requirements, breeding behavior, call and tadpole information are unknown. The following anuran species have been found sympatrically with the new species, at holotype locality: Limnonectes sp; Philautus larutensis; Sumaterana dabulescens Arifin, Smart, Hertwig, Smith, Iskandar, and Hass; at the paratype locality: Chalcorana chalconota; Leptophryne borbonica; Limnonectes kuhlii; L. macrodon; Limnonectes sp.; Rhacophorus catamitus; Philautus sp. and Pulchrana fantastica Arifin, Cahyadi, Smart, Jankowski, and Haas.

Megophrys selatanensis sp. nov. (Figs. 2 A−F, 3A−E) Holotype MZB. Amph 22411 (field number ENS 14208; GenBank accession no. MT 710704; Figs. 2A–B, 3A–E; Megophrys sp. south in Fig. 1 and Table 1), an adult female collected from the eastern side of Bukit Barisan Mountain range in southern Sumatra, Lampung Province, Tanggamus Regency, Ulubelu District, Ngarip (5.280170°S, 104.557240°E, 1439 m a.s.l., Fig. 6), by Elijah Wostl, Kyle A. O’Connell and Ahmad Muammar Kadafi at 2040 h on 10 June 2013. Paratype UTA A-66176 (formerly MZB. Amph 25988, field number ENS 17384; GenBank accession no. MT 710705; Fig. 2 C−D), subadult male collected at 1959 h on 8 July 2015 from the eastern side of Bukit Barisan Mountain range in southern Sumatra, Sumatra Selatan Province, Muara Enim Regency, Semendo Darat Ulu District, Segamit, near Gunung Patah (4.219890°S, 103.471250°E, 1624 m a.s.l., Fig. 6) by Elijah Wostl, Eric N. Smith, and Farid Akhsani. Referred specimens MZB. Amph 32593 (field number ENS 14202; GenBank accession no. MT 710706; Fig. 2F) a juvenile male (SVLh 16.08 mm), same locality and collector as holotype; UTA A-66177 (field number ENS 17403; GenBank accession no. MT 710707; Fig. 2E) a juvenile male (SVLh 23.5 mm), same locality and collectors as paratype. Etymology. The specific name selatanensis is derived from the Indonesian word Selatan (=south), as the new species exhibits a southern distribution within Sumatra and the Latin suffix – ensis meaning from that place. Suggested English common name. South-Sumatran Horned-Frog Suggested Indonesian name. Katak-tanduk Sumatra-selatan. Diagnosis. The new species was assigned to the genus Megophrys based on the combination of the following morphological characters, as defined by Kuhl and van Hasselt (1822) and Delorme et al. (2006): (1) pointed snout profile bearing a pointed projection, protruding laterally beyond the lower jaw; (2) broad and flattened eyelid with a palpebral projection; (3) possession of a broad and depressed head; (4) conical spine at the corner of the mouth; (5) vertical pupil; (6) and presence of maxillary and vomerine teeth. Megophrys selatanensis sp. nov. can be diagnosed from geographically proximate congeners from the Sunda Shelf and the Philippines by the following combination of morphological characters: large body, stocky (SVLh 79.3 mm in adult female); snout acute with short rostral appendage (RSAL 0.3% in adult female); relatively short triangular palpebral projection with acute tip (EHL 21.3% UEWh in adult female); head wider than long (HW 126.4% HLh in adult female); tympanum distinct, vertically elongated (TDH 52.4% TDV in adult female); vomerine teeth present; a pair of dorsolateral folds extending from shoulder above axilla to groin; dorsal skin smooth with low and dense tuberculation; ventral skin smooth from throat to belly; short thigh (RTL 40.3% in adult female), foot nearly as long as the thigh (FL 99.4% TL in adult female); tibiotarsal articulation reaching posterior corner of eye; toes webbed at base. Description of holotype (measurements in mm). Adult female, large body size (SVLh 79.3; SVL 79.1) and habitus stocky; head depressed and broad, wider (HW 38.1, 48.0% SVLh) than long (HL 30.6, 38.6% SVLh); snout short (SL 9.4, 11.9% SVLh), pointed at tip, acute, with very short rostral appendage (SAL 0.3% SVLh), laterally protruding and projecting beyond lower jaw; nostril positions laterally, closer to tip than to snout; eye positioned laterally, large, over three and a half times horizontal diameter of tympanum (ED 9.9, 365.1% TDH), eye diameter (ED 9.9, 12.5% SVLh) as long as snout-horn length (SLh 9.8, 12.3% SVLh), about twice nostril-eye length (ED 210.6% NEL), pupil vertical elliptical; canthus rostralis with sharp and angular ridge, lore sloping and concave; internarial distance (IND 7.6, 9.6% SVLh) more than half of interorbital distance (IND 66.7% IOD); palpebral projection length about 0.2 times of total upper eyelid width (EHL 1.8, 21.3% UEWh), tip acute, surface smooth; upper eyelid edges with scattered small and low conical tubercles; tympanum distinct, smooth, oval, elongated vertically (TDH 2.7, 3.4% SVLh; TDV 5.2, 6.5% SVLh; TDH 52.4% TDV); angular supratympanic fold, distinct, extending from behind eye, curving down around upper border of tympanum and ending above axilla; white conical tubercles present behind supratympanic fold and front of axilla; spinous gland on the corner of mouth on jaw angle; single row of maxillary teeth present; vomerine teeth present in two widely separated groups, at the level of posterior borders of choana; tongue lanceolate, notched posteriorly, without papillae. Forelimb slender, short, hand length about half of arm length (HAL 21.3, 55.9% LAL), lower and upper arms slender; fingers moderately slender with rounded and swollen tips, unwebbed, lacking lateral fringes, finger length formula II=IV Hindlimb slender, moderately long (HLL 106.8, 134.7% SVLh), thigh (FML 36.7, 46.3% SVLh) slightly longer than tibia (TL 32.0, 40.1% SVLh), about twice of tarsus length (TSL 17.6, 22.2% SVLh), and nearly as long as foot (FL 31.8, 40.1% SVLh); toe length formula I Dorsal skin smooth, with low and dense tubercles, mostly on forelimbs, hindlimbs, and flanks; a pair of distinct dorsolateral folds, one on each side, extending from shoulder, above axilla to groin; vent without dorsal dermal accessory extension; three transverse folds on lower arm; five transverse folds on hindlimb, three on thigh and two on tibia; limbs skin laterally smooth, with small tubercles; ventral skin smooth, wrinkled in preservative; pair of white conical pectoral glands at base of axilla; a white conical femoral gland at mid flanks of posterior thigh. Coloration. In life (Fig. 2A–B): pupil dark, iris golden-brown; base colour of dorsum uniform light brown; dorsal surface of forearm, hand, and hindlimbs slightly lighter than dorsum; upper limbs with dark brown transverse folds, three on forearm, three on thigh, and two on tarsus, fingers with dark brown crossbars, one on first and second fingers, two on third and fourth fingers; knee with irregular dark spots; dorsal surface and lateral sides of head light brown, with dark spots on upper and lower lips, from jaw angle to tip of snout; skin around eyes dark brown; flanks light brown, brighter than dorsum, unmarked; ventral surface light brown, with darker throat and chest; longitudinal markings over throat to pectoral region and dark blotches on belly; light brown blotches on ventral of limbs, with dark brown pattern near the joint between hand and lower arm. In preservative, dorsal surface darker, lateral and ventral surfaces fading to whitish, but pattern remains (Fig. 3A–E). Variation. Type and referred specimens are morphometrically similar, despite large ontogenetic discrepancy between the adult female holotype (MZB. Amph 22411), subadult male paratype (UTA A-66176), largest juvenile referred specimen (UTA A-66177), and smaller juvenile referred specimen (MZB. Amph 32593). The adult specimen has slightly shorter rostral and palpebral projections, a shorter head, and shorter limbs, as compared to the subadult and juvenile specimens. The tibiotarsal joint, when the hindlimbs are adpressed forward, reaches the posterior corner of the eye in the adult and subadult, but only reaches the mid-eye in the juvenile. The morphometric variation is given in Table 2. In coloration, the dorsum of the subadult paratype (UTA A-66176, Fig. 2B) and the referred specimen (UTA A-66177, Fig. 2E) is brighter than that of the holotype, this is darker in the smaller referred specimen (MZB Amph 32593, Fig. 2F). An irregular brighter brown pattern below the canthus rostralis is present in the paratype and the referred specimens but, it is absent in the holotype (Fig. 2A, C, E–F). The paratype and referred specimens have a dark inverted triangle pattern on the parietal-orbital-scapular region. The paratype has orange-reddish coloration on the throat, chest, and ventral surface of the fingers and toes (Fig 2C, D). The juvenile specimens (MZB Amph 32593 and UTA A-66177) have darker brown throats and chests. Comparisons. Megophrys selatanensis sp. nov. is morphologically most similar to M. montana than to other geographically related congeners. Megophrys selatanensis sp. nov. has elongated dorsolateral folds that extend from the parietoscapular region to the groin, type I, as M. montana, M. parallela, M. lancip, M. nasuta, M. kalimantanensis Munir, Hamidy, Matsui, Iskandar, Sidik, and Shimada, and M. stejnegeri Taylor. Megophrys selatanensis sp. nov. differs from M. montana by having a shorter rostral appendage—SAL 0.3−0.6% SVLh (vs. SAL reaching 3.5% SVLh: Munir et al., 2018), shorter palpebral projection—EHL 21.3–24.3% UEWh (vs. reaching 48.4% UEWh: Munir et al., 2018), shorter ratio of foot to thigh—FL 96.6−99.4% TL (vs. FL 106.2−115.6% TL: Munir et al., 2018), and toes webbed only at base (vs. more developed web on second to fifth toes: Munir et al., 2018). Megophrys selatanensis sp. nov. differs from M. parallela by having elongated dorsolateral folds reaching the groin (vs. only reaching two-thirds of trunk), females being large—known adult female 79.1 mm SVL (vs. SVL ≤ 58.3 mm: Munir et al., 2018), a rostral appendage present (vs. absent: Inger and Iskandar, 2005; Munir et al., 2018), shorter ratio of foot to thigh—FL 96.6–99.4% TL (vs. FL 105.4–112.5% TL: Munir et al., 2018). The new species differs from M. lancip by having a less developed rostral appendage—SAL 0.3−0.6% SVLh (vs. SAL 1.4–4.1% SVLh: Munir et al., 2018), a shorter head length—HLh 38.0–38.5% SVLh (vs. HLh 42.1–44.6% SVLh: Munir et al., 2018), slightly wider head ratio to its length—HW 119.3–126.4% HLh (vs. HW 103.2–116.0% HLh: Munir et al., 2018), relatively shorter ratio of feet to thigh—FL 96.6–99.4%TL (vs. FL 102.5–109.8% TL: Munir et al., 2018), toes webbed only at base (vs. rather developed on third, fourth and fifth toes: Munir et al., 2018). ......continued on the next page Megophrys selatanensis sp. nov. differs from M. nasuta by the absence of additional lateral folds on flanks (vs. presence: Munir et al., 2018, 2019), shorter acute rostral appendage—SAL 0.3−0.6% SVLh (vs. acuminate rostral appendage, SAL 1.8−9.2% SVLh: Munir et al., 2019), shorter acute palpebral projection—EHL 2.2−3.4% SVLh (vs. acuminate palpebral projection, EHL 4.4−13.0% SVLh: Munir et al., 2019), shorter snout to nostril length—SNLh 3.7−5.1% SVLh (vs. SNLh 5.1−13.8% SVLh: Munir et al., 2019), shorter head length—HLh 38.0−40.5% SVLh (vs. HLh 39.5−51.7.% SVLh: Munir et al., 2019), smaller ratio of palpebral projection to upper eyelid width—EHL 21.3−25.7% UEWh (vs. EHL 32.7−61.4% UEWh: Munir et al., 2019), and relatively wider head—HW 119.3−126.4% HLh (vs. HW 92.9−119.5% HLh: Munir et al., 2019). From M. kalimantanensis, the new species differs by the absence of additional lateral folds on flanks (vs. presence: Munir et al., 2019), female being smaller—known adult female 79.1 mm SVL (vs. 109.1−116.4 mm SVL: Munir et al., 2019), adpressed tibiotarsal articulation reaching posterior corner of eye in female (vs. posterior of tympanum, in female: Munir et al., 2019). From M. stejnegeri Taylor, Megophrys selatanensis sp. nov. differs by having the dorsolateral folds extending to the groin (vs. maximum of two-thirds length of trunk: Inger, 1954), female being slightly larger—known adult female 79.1 mm SVL (vs. SVL 57.0– 77.8 mm: Taylor, 1920; Inger 1954), a rostral appendage present (vs. absent: Taylor, 1920; Inger, 1954), vomerine teeth present (vs. absent: Taylor, 1920; Inger, 1954), tympanum vertically elongated (vs. slightly rounded, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016), and smooth skin on the temporal region (vs. tuberculate, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016). Megophrys selatanensis sp. nov. is distinguished from M. kobayashii Malkmus and Matsui by having type I dorsolateral folds (vs. dorsolateral folds extended from parietoscapular region to mid-body, type II, see Table 3), additional lateral folds on flanks absent (vs. present: Malkmus and Matsui, 1997; Munir et al., 2018), female being smaller—known adult female 79.1 mm SVL (vs. SVL 99.0–109.0 mm: Malkmus and Matsui, 1997; Inger and Stuebing, 2005), a rostral appendage present (vs. absent: Malkmus and Matsui, 1997), and smaller and fewer tubercles on the body flanks (vs. larger and numerous: Malkmus and Matsui, 1997). From M. ligayae Taylor, the new species differs in having type I dorsolateral folds (vs. type II: Taylor, 1920; Inger, 1954; Munir et al., 2018), no additional lateral folds on flanks (vs. present: Taylor, 1920; Inger, 1954; Munir et al., 2018), female being smaller—known adult female 79.1 mm SVL (vs. SVL 90.0 mm: Inger, 1954), tibiotarsal articulation reaching posterior corner of eye (vs. tympanum: Taylor, 1920), and toes webbed only at base (vs. rather developed web on third, fourth, and fifth toes, see figure 37C Diesmos et al., 2015). From M. edwardinae Inger, the new species differs by having type I dorsolateral folds (vs. absent: Inger, 1989), a rostral appendage present (vs. absent: Inger, 1989), vomerine teeth present (vs. absent: Inger, 1989), and smooth dorsal skin surface with low and dense tubercles (vs. dorsal surface with scattered round and elongated tubercles: Inger, 1989). Megophrys selatanensis sp. nov. can be distinguished from M. baluensis Boulenger by having type I dorsolateral folds (vs. dorsolateral folds formed by a series of elongated tubercle, type III: Inger and Stuebing, 2005: Munir et al., 2018), female being slightly larger—known adult female 79.1 mm SVL (vs. SVL ≤ 70 mm: Inger, 1966; Inger et al., 1995), a rostral appendage present (vs. absent: Boulenger, 1899; Inger, 1966; Inger et al., 1995), having a triangular palpebral projection (vs. small palpebral projection like tubercle: Inger, 1989; Inger et al., 1995), having smaller and fewer tubercles on body flanks (vs. larger and numerous: Inger et al., 2017), and adpressed tibiotarsal articulation reaching posterior of eye (vs. shoulder: Boulenger, 1899). Megophrys selatanensis sp. nov. differs from M. dringi Inger, Stuebing, and Tan, M. aceras Boulenger, and M. longipes Boulenger by the absence of Y, X,>–et al., 1995, see Table 3). Furthermore, from M. dringi the new species being larger in female—known adult female 79.1 mm SVL (vs. SVL 55 mm: Inger et al., 1995; Inger and Stuebing, 2005), having a stocky body (vs. slender: Inger et al., 1995), a rostral appendage present (vs. absent: Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger et al., 1995), vomerine teeth present (vs. absent: Inger et al., 1995), distinct tympanum (vs. partially obscured: Inger et al., 1995). The new species differs from M. aceras by having a stocky body (vs. slender: Boulenger, 1903), a rostral appendage present (vs. absent: Boulenger, 1903), tibiotarsal articulation reaching to posterior corner of eye (vs. shoulder, angle of jaws or temporal area: Taylor, 1962), and toes webbed only at base (vs. rather possessing developed web between third, fourth, and fifth toes: Taylor, 1962; see figure 5 in Munir et al., 2018). From M. longipes, the new species differs being larger in female—known adult female 79.1 mm SVL (vs. SVL 49.0–65.0 mm: Inger et al., 1995), having a stocky body (vs. slender: Boulenger, 1885), a rostral appendage present (vs. absent: Boulenger, 1885), having a triangular palpebral projection (vs. small like tubercle: Boulenger, 1885; Taylor, 1962), shorter thigh—TL 0.41−0.44 SVL (vs. TL 0.54–0.65 SVL: Inger et al., 1995; Inger and Iskandar, 2005) and tibiotarsal articulation reaching posterior corner of eye (vs. far beyond tip of snout: Boulenger, 1885; Taylor, 1962). Distribution and natural history. The holotype and one referred specimen (MZB Amph 32593) were collected in leaf litter on the edge of an old secondary forest in proximity to a coffee plantation and above a geothermal station in the province of Lampung, while the paratype (UTA A-66176) and the other referred specimen (UTA A- 66177) were collected from an old secondary forest in the green corridor between the Gunung Patah and Gunung Agung. These locations are about 170 km apart along the eastern slope of the southern Barisan range of Sumatra. The tadpoles (ENS 14306 and ENS 14737) were collected from a small rocky stream in the old secondary forest near Lake Ranau, Lampung, 1108–1487 m a.s.l. (description of the tadpole of this new species is pending as we are preparing another manuscript to present detailed description of Sumatran Megophrys tadpoles). Most of the samples of M. selatanensis sp. nov. were collected from a small patched forest area in the eastern Bukit Barisan Selatan Mountain ranges, which is threatened by high deforestation. The exact distribution, population, breeding behavior, call, and other ecological information are unknown. The following anuran species have been found sympatrically with the new species, at Ngarip, Lampung: Indosylvirana nicobariensis (Stoliczka); Chalcorana chalconota (Schlegel); Wijayarana sumatrana (Yang); Hylarana erythraea (Schlegel); Ingerophrynus biporcatus (Gravenhorst); Leptophryne borbonica (Tschudi); Limnonectes kuhlii (Tschudi); L. macrodon (Duméril and Bibron); Limnonectes sp.; Microhyla gadjahmadai Atmaja, Hamidy, Arisuryanti, Matsui, Smith; M. palmipes Boulenger; Pelophryne sp.; Philautus larutensis (Boulenger); P. polymorphus Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; P. thamyridion Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; Phrynoides asper (Gravenhorst); Polypedates leucomystax (Gravenhorst); Rhacophorus achantharrhena

Megophrys acehensis sp. nov. (Figs. 4 A���D, 5A���E) Holotype MZB. Amph 26098 (field number ENS 18664; GenBank accession no. MT 710708; Figs. 4 A���B, 5A���E; Megophrys sp. north in Fig. 1 and Table 1), adult male collected at 2330 h on 5 August 2015 from Aceh Province, Aceh Tengah Regency, Linge District, Kute Robel (4.506444 ��N, 96.860750 ��E, 1638 m a.s.l), by Elijah Wostl, Ilham Fonna, and Muhammad Iksan (Fig. 6). Paratype UTA A���66178 (field number ENS 21030; GenBank accession no. MT 710709; Fig. 4C���D), a sub adult male collected on at 1856 h on 1 June 2016 from Aceh Province, Pidie Regency, Geumpang District, UPT V Geumpang (4.854540 ��N, 96.216540 ��E, 1086 m a.s.l) by Michael B. Harvey and Eric N. Smith (Fig. 6). Etymology. The specific name acehensis is derived from the province of Aceh in northern Sumatra and the Latin suffix��� ensis meaning from that place. Suggested English common name. Aceh Horned Frog Suggested Indonesian name. Katak���tanduk Aceh Diagnosis. The new species was assigned to the genus Megophrys based on the combination of the following morphological characters, as defined by Kuhl and van Hasselt (1822) and Delorme et al. (2006): (1) pointed snout profile, bearing a pointed projection, protruding laterally beyond the lower jaw; (2) broad and flattened eyelid with palpebral projection; (3) possession of a broad and depressed head; (4) conical spine at the corner of mouth; (5) vertical pupil; (6) presence of maxillary and vomerine teeth. Megophrys acehensis sp. nov. can be diagnosed from its geographically relevant congeners in the Sunda Shelf and the Philippines by the following combination of characters: medium body size, stocky (SVLh 38.1 mm in adult male); snout pointed with a short, acute rostral appendage (RSAL 1.3% in adult male); relatively short triangular palpebral projection with acute tip (EHL 25.1% UEWh in adult male); head length relatively short (RHLh 37.2% in adult male); head wider than its long (HW 113.6% HLh in adult male); tympanum distinct, about one-third of eye diameter (TDH 35.6% ED in adult male), nearly rounded (TDH 85.9 % TDV in adult male); vomerine teeth present; a pair of dorsolateral folds, extending from shoulder, above axilla to groin; dorsal and lateral skin tuberculate, shagreened on the throat to belly; short lower arm (RLAL 45.8% in adult male); foot nearly as long as thigh (FL 96.6% TL in adult male); tibiotarsal articulation reaching posterior of eye; toe webbing absent. Description of holotype (measurements in mm). Adult male, medium body size (SVLh 38.1, SVL 37.4) and habitus stocky; head depressed and broad, wider (HW 16.4, 42.9% SVLh) than long (HL 13.3, 34.9% SVLh); snout short (SL 4.2, 11.1% SVLh), pointed at tip with acute short rostral appendage (SAL 0.7, 1.8% SVLh), laterally protruding and projecting beyond lower jaw; nostril positioned laterally, near to snout than to eye; eye positioned laterally, large, nearly three times of tympanum horizontal diameter (ED 5.5, 280.5% TDH), eye diameter slightly wider (ED 5.5, 14.4% SVLh) than snout���horn length (SLh 5.4, 14.2% SVLh), about two and three-quarter times of nostril���eye length (ED 275% NEL), pupil vertical elliptical; canthus rostralis with sharp, angular ridge, lore sloping and concave; internarial distance (IND 4.0, 10.4% SVLh) about two-thirds of interorbital distance (IND 68.4% IOD); palpebral projection length about one-quarter of total upper eyelid width (EHL 1.2, 25.1% UEWh), tip acute, surface smooth and scattered with small and low tubercles; tympanum distinct, smooth, oval, slightly rounded (TDV 2.3, 6.0% SVLh; TDH 2.0, 5.1% SVLh; TDH 85.9% TDV); angular supratympanic fold, distinct, widened anteriorly, narrowed posteriorly, extending from behind eye, curving down around upper border of tympanum and ending above axilla; white conical tubercles behind the supratympanic fold and anterior to axilla; spinous gland on corner of mouth on jaw angle; single row of maxillary teeth present; vomerine teeth in two widely separated groups, at level posterior borders of choana; tongue lanceolate, notched posteriorly, without papillae; median subgular vocal sac present, having slit���like opening on each side of jaw commissures. * Fold forming Y, X or H on the parietoscapular region to the level of axilla; ** Dorsolateral fold shape: dorsolateral folds are elongated and extend from the parietoscapular region to the groin (Type I); dorsolateral folds extend from the central of parietoscapular region to mid-body (Type II); multiple dorsolateral folds - at least three or four - and they are discontinuous, formed by a series of elongated tubercles (Type III); dorsolateral folds are elongated and curve from the axillary region towards (and reaching) the posterior dorsal margin of tympanum (Type IV). Forelimb slender and short, hand length about half of arm length (HAL 9.5, 54.3% LAL), lower arm proximally enlarged, wider than upper arm; fingers moderately slender, with rounded and swollen tips, unwebbed and lacking of lateral fringes; finger length formula IColoration. In life (Fig. 4A���B, 5A���B), pupil dark, iris golden���brown, base of dorsum immaculate dark brown with indistinct dark brown inverted triangle pattern on parieto-orbital to scapular region and inverted ���V��� like pattern on back, positioned closer to vent than to parietal region; dorsal surface of head, forelimb and hindlimb darker than lateral body; dark brown transverse folds present on dorsal surface of forearm, thigh and tibia, two on the forearm, three on thigh and three on tibia, fingers with dark brown crossbars, one on first and second fingers, two on third and fourth fingers; dorsomedial surfaces of first and second fingers with dark brown microspinular nuptial pads; knee with irregular dark brown spots; dorsal surface and lateral sides of head dark brown, irregular lighter brown pattern below canthus rostralis, light brown on underside of eyelids; body flanks dark brown and unmarked, lighter than dorsum; forelimb flanks dark brown, as dorsum; hindlimb flanks cream with dark brown blotches; ventral surface from throat to chest dark brown with heavy dark longitudinal marking; belly and underside of fore- and hindlimbs light cream with dark brown blotches; in preservative, pattern remains, but dorsal, lateral, and ventral surfaces are darker (Fig. 5C). Variation. The two type specimens are morphometrically similar. The holotype has slightly longer rostral appendage, head, and snout-nostril and snout lengths, but a slightly shorter palpebral projection, and a thicker and narrower head. Morphometric variation is shown in Table 2. The transverse fold in the occipital region varies in degree of development, being distinct on the holotype but indistinct on the paratype. The dorsal coloration in life is dark brown on the holotype and orange-brown on the paratype. The ventral color of the two specimens is marbled with brown in both specimens but, the background color of the holotype is cream anteriorly and light grey posteriorly while that of the paratype is dark orange anteriorly and whitish posteriorly (Fig. 4 A���D). Comparisons. Megophrys acehensis sp. nov. differs from its most similar congener, M. parallela by having dorsolateral folds extending to the groin (vs. maximum of two-thirds length of trunk: Inger and Iskandar, 2005; Munir et al., 2018), a rostral appendage present (vs. absent: Inger and Iskandar, 2005; Munir et al., 2018), shagreened ventral surface from throat to belly (vs. smooth: Inger and Iskandar, 2005; Munir et al., 2018), and a relatively shorter foot to thigh ratio���FL 92.1���96.6% TL (vs. 105.4���112.5% TL: Munir et al., 2018). From M. montana, the new species differs by having tuberculate dorsal skin surface (vs. smooth: Munir et al., 2018), shagreened skin on throat (vs. smooth: Munir et al., 2018), a relatively short foot to thigh ratio���FL 92.1���96.6% TL (vs. FL 106.2���115.6% TL: Munir et al., 2018), and absent toe webbing (vs. rather developed webbing from second to fifth toes: Munir et al., 2018). From Megophrys selatanensis sp. nov., Megophrys acehensis sp. nov. differs by having a longer rostral appendage���SAL 1.3���1.8% SVLh (vs. SAL 0.3���0.6% SVLh), a slightly longer palpebral projection ���EHL 25.1���28.4% UEWh (vs. EHL 21.3���24.3% UEWh), slightly narrower ratio of head width to its length���HW 114.1���115.3% HLh (vs. HW 119.3���126.4% HLh), wider ratio of tympanum to eye diameter���TDH 35.6���37.5% ED (vs. TDH 27.3���28.5% ED), tympanum nearly rounded���TDH 85.9���93.8 TDV (vs. vertically elongated, TDH 46.5���61.9% TDV), slightly shorter lower arms���LAL 44.8���45.8% SVLh (vs. LAL 48.0���49.2% SVLh), tuberculate dorsal skin surface (vs. smooth), shagreened skin on throat (vs. smooth), and absent toe webbing (vs. toes webbed at base). Megophrys acehensis sp. nov. differs from M. lancip by males having a shorter rostral appendage���SAL 1.3���1.8% SVLh (vs. SAL 2.9���3.1% SVLh in males, but overlapped with females SAL 1.4���4.1% SVLh: Munir et al., 2018), shorter head length���HLh 37.2���37.8% SVLh (vs. HLh 42.1���44.6% SVLh; Munir et al., 2018), wider head in males���HW 114.1���115.3% HLh (vs. HW 103.2���105.8% HLh in males, but overlapped with females HW 110.6���116.0% HLh: Munir et al., 2018), wider tympanum relative to eye diameter���TDH 35.6���37.5% ED (vs. TDH 25.7���28.0 ED: Munir et al., 2018), tympanum nearly rounded���TDH 85.9���93.8 TDV (vs. vertically elongated, TDH 61.1���69.1% TDV), relatively short foot to thigh ratio���FL 92.1���96.6% TL (vs. FL 102.5���109.8% TL; Munir et al., 2018), and toe webbing absent (vs. rather developed on first to fifth toes, see Figs. 4D, I, 5B in Munir et al., 2018). Megophrys acehensis sp. nov. differs from M. nasuta by the absence of additional lateral flank folds (vs. presence: Munir et al., 2018, 2019), males being smaller���known adult male 37.4 mm SVL (vs. SVL 66.0��� 93.4 mm: Munir et al., 2019), shorter acute rostral appendage���SAL 1.3���1.8% SVLh (vs. acuminate rostral appendage, SAL 1.8���9.2% SVLh: Munir et al., 2019), and shorter acute palpebral projection���EHL 25.1���28.4% UEWh (vs. acuminate palpebral projection, EHL 32.7���61.4% UEWh: Munir et al., 2019). From M. kalimantanensis, the new species differs by the absence of additional flank folds (vs. present: Munir et al., 2019), smaller body size in males���known adult male 37.4 mm SVL (vs. SVL 64.3���100.6 mm: Munir et al., 2019), slightly longer rostral appendage���SAL 1.3���1.8% SVLh (vs. SAL 0.1���0.8% SVLh: Munir et al., 2019), and shorter palpebral projection���EHL 25.1���28.4% UEWh (vs. 30.9���53.2% UEWh: Munir et al., 2019). From M. stejnegeri, the new species differs by having dorsolateral folds extending to the groin (vs. maximum of two-thirds the length of the trunk: Inger 1954), males being smaller���known adult male 37.4 mm SVL (vs. adult male 40.6���60.0 mm: Taylor, 1920; Inger, 1954), vomerine teeth present (vs. absent: Taylor, 1920; Inger et al., 1954), and smooth skin on the posterior of tympanum (vs. tuberculate, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016). Megophrys acehensis sp. nov. can be distinguished from M. kobayashii by having type I dorsolateral folds (vs. type II, see Table 3), absent additional lateral flank folds (vs. present: Malkmus and Matsui, 1997; Munir et al., 2018), males being smaller���known adult male 37.4 mm SVL (vs. 93.0���101.0 mm: Inger and Stuebing, 2005), rostral appendage present (vs. absent: Malkmus and Matsui, 1997), and smaller and fewer tubercles on flanks (vs. larger and numerous: Malkmus and Matsui, 1997). From M. ligayae, the new species differs by having type I dorsolateral folds, extended to the groin (vs. type II, reaching a maximum of two-thirds of trunk: Taylor, 1920; Inger, 1954; Munir et al., 2018), absent additional lateral folds (vs. present: Taylor, 1920; Inger, 1954; Munir et al., 2018), males being smaller���known adult male 37.4 mm SVL (vs. SVL 60.4���69.0 mm: Taylor, 1920; Inger, 1954), and toe webbing absent (vs. rather developed web on third, fourth, and fifth toes: Diesmos et al., 2015). Megophrys acehensis sp. nov. differs from M. edwardinae by having type I dorsolateral folds (vs. absent: Inger 1989), a rostral appendage present (vs. absent: Inger 1989), vomerine teeth present (vs. absent: Inger 1989), and slightly shorter thighs���TL 40.4���41.8% SVLh (vs. TL 45.0���50.0% SVL: Inger 1989). Megophrys acehensis sp. nov. can be distinguished from M. baluensis by having type I dorsolateral folds (vs. type III: Munir et al., 2018), males being slightly smaller���known adult male 37.4 mm SVL (vs. SVL 41.0���45.0 mm: Inger et al., 1995), a rostral appendage present (vs. absent: Boulenger, 1899; Inger, 1966; Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger, 1989; Inger et al., 1995), having smaller and fewer tubercles on body flanks (vs. larger and numerous: Inger et al., 2017), and adpressed tibiotarsal articulation reaching posterior of eye (vs. shoulder: Boulenger, 1899). Megophrys acehensis sp. nov. can be distinguished from M. dringi, M. aceras and M. longipes by the absence of Y, X,>���et al., 1995, see Table 3). Furthermore, from M. dringi, the new species being smaller in male���known adult male 37.4 mm SVL (vs. SVL 43.0���47.0 mm: Inger et al., 1995), having a stocky body (vs. slender: Inger et al., 1995), a rostral appendage present (vs. absent: Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger et al., 1995), vomerine teeth present (vs. absent: Inger et al., 1995), distinct tympanum (vs. partially obscured: Inger et al., 1995). From M. aceras, the new species differs by having smaller body size in male���known adult male 37.4 mm SVL (vs. SVL 48.0��� 62.4 mm: Inger and Iskandar, 2005), a stocky body (vs. slender: Boulenger, 1903), rostral appendage present (vs. absent: Boulenger, 1903), tibiotarsal articulation reaching to posterior corner of eye (vs. shoulder, angle of jaws or temporal area: Taylor, 1962), toes web absent (vs. rather developed web on third, fourth, and fifth toes: Taylor, 1962, see figure 5 in Munir et al., 2018). From M. longipes, the new species differs by having a smaller body size in male���known adult male 37.4 mm SVL (vs. SVL 38.9���45.2 mm: Inger and Iskandar, 2005), having a stocky body (vs. slender: Boulenger, 1885), a rostral appendage present (vs. absent: Boulenger, 1885), having a triangular palpebral projection (vs. small like tubercle: Boulenger, 1885; Taylor, 1962), shorter thigh���TL 0.41���0.42 SVL (vs. TL 0.55���0.60 SVL: Inger and Iskandar, 2005) and tibiotarsal articulation reaching posterior corner of eye (vs. far beyond tip of snout; Boulenger 1885). Distribution and Natural History. The holotype of Megophrys acehensis sp. nov. was collected on leaf litter in a sloping area at the edge of a primary forest near a stream, while the paratype was collected from leaf litter in a palm oil plantation near the edge of an old secondary forest. Landslides, new road development, and monoculture forests have become major threats at the holotype locality, while the threats at the paratype locality were the land use changes and water pollution from palm oil fields. The precise distribution, population, habitat requirements, breeding behavior, call and tadpole information are unknown. The following anuran species have been found sympatrically with the new species, at holotype locality: Limnonectes sp; Philautus larutensis; Sumaterana dabulescens Arifin, Smart, Hertwig, Smith, Iskandar, and Hass; at the paratype locality: Chalcorana chalconota; Leptophryne borbonica; Limnonectes kuhlii; L. macrodon; Limnonectes sp.; Rhacophorus catamitus; Philautus sp. and Pulchrana fantastica Arifin, Cahyadi, Smart, Jankowski, and Haas., Published as part of Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir & Smith, Eric N., 2021, Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia, pp. 503-529 in Zootaxa 5057 (4) on pages 514-520, DOI: 10.11646/zootaxa.5057.4.3, http://zenodo.org/record/5598859, {"references":["Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24 (1 - 4), 6 - 21.","Munir, M., Hamidy, A., Farajallah, A. & Smith, E. N. (2018) A new Megophrys Kuhl and van Hasselt (Amphibia: Megophryidae) from southwestern Sumatra, Indonesia. Zootaxa, 4442 (3), 389 - 412. https: // doi. org / 10.11646 / zootaxa. 4442.3.3","Inger, R. F. & Iskandar, D. T. (2005) A collection of amphibians from West Sumatra, with description of new species of Megophrys (Amphibia: Anura). The Raffles Bulletin of Zoology, 53 (1), 133 - 142.","Munir, M., Hamidy, A., Matsui, M., Iskandar, D. T., Sidik, I. & Shimada, T. (2019) A new species of Megophrys Kuhl & van Hasselt (Amphibia: Megophryidae) from Borneo allied to M. nasuta (Schlegel, 1858). Zootaxa, 4679 (1), 1 - 24. https: // doi. org / 10.11646 / zootaxa. 4679.1.1","Inger, R. F. (1954) The systematics and zoogeography of Philippine Amphibia. Fieldiana Zoology, 33 (4), 183 - 531. https: // doi. org / 10.5962 / bhl. title. 5571","Taylor, E. H. (1920) Philippine Amphibia. The Philippine Journal of Science, 16 (3), 213 - 359. https: // doi. org / 10.5962 / bhl. part. 4751","Diesmos, A. C., Watters, J. L., Huron, N. A., Davis, D. R., Alcala, A. C., Crombie R. I., Afuang, L. E., Gee-das, G., Sison, R. V., Sanguila, M. B., Penrod, M. L., Labonte, M. J., Davey C. S., Leone E. A., Diesmos M. L., Sy, M. Y., Welton, L. J., Brown R. M. & Siler C. D. (2015) Amphibians of the Philippines, Part I: Checklist of the Species. Proceedings of the California Academy of Science, 62 (20), 457 - 539.","Sanguila, M. B., Cobb, K. A, Siller C. D., Diesmos, A. C., Alcala A. C. & Brown R. M. (2016) The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands. ZooKeys, 624, 1 - 32. https: // doi. org / 10.3897 / zookeys. 624.9814","Malkmus, R. & Matsui, M. (1997) Megophrys kobayashii, ein neuer pelobatider frosch vom Mount Kinabalu, Sauria, 19, 31 - 37.","Inger, R. F. & Stuebing, R. B. (2005) A field guide to the frogs of Borneo. 2 nd Edition. Natural History Publication Borneo Sdn. Bhd. Kota Kinabalu, Sabah, 201 pp.","Inger, R. F. (1989) Four new species of frogs from Borneo. Malayan Nature Journal, 42, 229 - 243.","Inger, R. F., Stuebing, R. B. & Tan, F. L. (1995) New species and new records of anurans from Borneo. The Raffles Bulletin of Zoology, 43 (1), 115 - 131.","Boulenger, G. A. (1899) Description of three new reptiles and a new batrachian from Mount Kinabalu, North Borneo. Annals and Magazine of Natural History, 7 (4), 451 - 454. https: // doi. org / 10.1080 / 00222939908678228","Inger, R. F. (1966) The systematic and zoogeography of the amphibians of Borneo. Fieldiana Zoology, 52, 1 - 402. https: // doi. org / 10.5962 / bhl. title. 3147","Inger, R. F., Stuebing, R. B., Grafe, T. U. & Dehling, J. M. (2017). A field guide to the Frogs of Borneo. Third edition. Natural History Publication Borneo. Kinabalu, 236 pp.","Boulenger, G. A. (1885) Description of a new frog of the genus Megalophrys. Proceedings of the scientific meeting of the Zoological Society of London, 53 (4), 850 - 850. https: // doi. org / 10.1111 / j. 1096 - 3642.1885. tb 02927. x","Boulenger, G. A. (1903) Report on the batrachians and reptiles. In: Annandale, N. & Robinson, H. C. (Eds.), Fasciculi Malayenses. Zoology. Part I. Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States 1901 - 1902. Longmans, Green & Co, London, pp. 131 - 176.","Taylor, E. H. (1962) The amphibian fauna of Thailand. The University of Kansas Science Bulletin, 43 (8), 265 - 599. https: // doi. org / 10.5962 / bhl. part. 13347"]}

Megophrys selatanensis sp. nov. (Figs. 2 A���F, 3A���E) Holotype MZB. Amph 22411 (field number ENS 14208; GenBank accession no. MT 710704; Figs. 2A���B, 3A���E; Megophrys sp. south in Fig. 1 and Table 1), an adult female collected from the eastern side of Bukit Barisan Mountain range in southern Sumatra, Lampung Province, Tanggamus Regency, Ulubelu District, Ngarip (5.280170��S, 104.557240��E, 1439 m a.s.l., Fig. 6), by Elijah Wostl, Kyle A. O���Connell and Ahmad Muammar Kadafi at 2040 h on 10 June 2013. Paratype UTA A-66176 (formerly MZB. Amph 25988, field number ENS 17384; GenBank accession no. MT 710705; Fig. 2 C���D), subadult male collected at 1959 h on 8 July 2015 from the eastern side of Bukit Barisan Mountain range in southern Sumatra, Sumatra Selatan Province, Muara Enim Regency, Semendo Darat Ulu District, Segamit, near Gunung Patah (4.219890��S, 103.471250��E, 1624 m a.s.l., Fig. 6) by Elijah Wostl, Eric N. Smith, and Farid Akhsani. Referred specimens MZB. Amph 32593 (field number ENS 14202; GenBank accession no. MT 710706; Fig. 2F) a juvenile male (SVLh 16.08 mm), same locality and collector as holotype; UTA A-66177 (field number ENS 17403; GenBank accession no. MT 710707; Fig. 2E) a juvenile male (SVLh 23.5 mm), same locality and collectors as paratype. Etymology. The specific name selatanensis is derived from the Indonesian word Selatan (=south), as the new species exhibits a southern distribution within Sumatra and the Latin suffix ��� ensis meaning from that place. Suggested English common name. South-Sumatran Horned-Frog Suggested Indonesian name. Katak-tanduk Sumatra-selatan. Diagnosis. The new species was assigned to the genus Megophrys based on the combination of the following morphological characters, as defined by Kuhl and van Hasselt (1822) and Delorme et al. (2006): (1) pointed snout profile bearing a pointed projection, protruding laterally beyond the lower jaw; (2) broad and flattened eyelid with a palpebral projection; (3) possession of a broad and depressed head; (4) conical spine at the corner of the mouth; (5) vertical pupil; (6) and presence of maxillary and vomerine teeth. Megophrys selatanensis sp. nov. can be diagnosed from geographically proximate congeners from the Sunda Shelf and the Philippines by the following combination of morphological characters: large body, stocky (SVLh 79.3 mm in adult female); snout acute with short rostral appendage (RSAL 0.3% in adult female); relatively short triangular palpebral projection with acute tip (EHL 21.3% UEWh in adult female); head wider than long (HW 126.4% HLh in adult female); tympanum distinct, vertically elongated (TDH 52.4% TDV in adult female); vomerine teeth present; a pair of dorsolateral folds extending from shoulder above axilla to groin; dorsal skin smooth with low and dense tuberculation; ventral skin smooth from throat to belly; short thigh (RTL 40.3% in adult female), foot nearly as long as the thigh (FL 99.4% TL in adult female); tibiotarsal articulation reaching posterior corner of eye; toes webbed at base. Description of holotype (measurements in mm). Adult female, large body size (SVLh 79.3; SVL 79.1) and habitus stocky; head depressed and broad, wider (HW 38.1, 48.0% SVLh) than long (HL 30.6, 38.6% SVLh); snout short (SL 9.4, 11.9% SVLh), pointed at tip, acute, with very short rostral appendage (SAL 0.3% SVLh), laterally protruding and projecting beyond lower jaw; nostril positions laterally, closer to tip than to snout; eye positioned laterally, large, over three and a half times horizontal diameter of tympanum (ED 9.9, 365.1% TDH), eye diameter (ED 9.9, 12.5% SVLh) as long as snout-horn length (SLh 9.8, 12.3% SVLh), about twice nostril-eye length (ED 210.6% NEL), pupil vertical elliptical; canthus rostralis with sharp and angular ridge, lore sloping and concave; internarial distance (IND 7.6, 9.6% SVLh) more than half of interorbital distance (IND 66.7% IOD); palpebral projection length about 0.2 times of total upper eyelid width (EHL 1.8, 21.3% UEWh), tip acute, surface smooth; upper eyelid edges with scattered small and low conical tubercles; tympanum distinct, smooth, oval, elongated vertically (TDH 2.7, 3.4% SVLh; TDV 5.2, 6.5% SVLh; TDH 52.4% TDV); angular supratympanic fold, distinct, extending from behind eye, curving down around upper border of tympanum and ending above axilla; white conical tubercles present behind supratympanic fold and front of axilla; spinous gland on the corner of mouth on jaw angle; single row of maxillary teeth present; vomerine teeth present in two widely separated groups, at the level of posterior borders of choana; tongue lanceolate, notched posteriorly, without papillae. Forelimb slender, short, hand length about half of arm length (HAL 21.3, 55.9% LAL), lower and upper arms slender; fingers moderately slender with rounded and swollen tips, unwebbed, lacking lateral fringes, finger length formula II=IVColoration. In life (Fig. 2A���B): pupil dark, iris golden-brown; base colour of dorsum uniform light brown; dorsal surface of forearm, hand, and hindlimbs slightly lighter than dorsum; upper limbs with dark brown transverse folds, three on forearm, three on thigh, and two on tarsus, fingers with dark brown crossbars, one on first and second fingers, two on third and fourth fingers; knee with irregular dark spots; dorsal surface and lateral sides of head light brown, with dark spots on upper and lower lips, from jaw angle to tip of snout; skin around eyes dark brown; flanks light brown, brighter than dorsum, unmarked; ventral surface light brown, with darker throat and chest; longitudinal markings over throat to pectoral region and dark blotches on belly; light brown blotches on ventral of limbs, with dark brown pattern near the joint between hand and lower arm. In preservative, dorsal surface darker, lateral and ventral surfaces fading to whitish, but pattern remains (Fig. 3A���E). Variation. Type and referred specimens are morphometrically similar, despite large ontogenetic discrepancy between the adult female holotype (MZB. Amph 22411), subadult male paratype (UTA A-66176), largest juvenile referred specimen (UTA A-66177), and smaller juvenile referred specimen (MZB. Amph 32593). The adult specimen has slightly shorter rostral and palpebral projections, a shorter head, and shorter limbs, as compared to the subadult and juvenile specimens. The tibiotarsal joint, when the hindlimbs are adpressed forward, reaches the posterior corner of the eye in the adult and subadult, but only reaches the mid-eye in the juvenile. The morphometric variation is given in Table 2. In coloration, the dorsum of the subadult paratype (UTA A-66176, Fig. 2B) and the referred specimen (UTA A-66177, Fig. 2E) is brighter than that of the holotype, this is darker in the smaller referred specimen (MZB Amph 32593, Fig. 2F). An irregular brighter brown pattern below the canthus rostralis is present in the paratype and the referred specimens but, it is absent in the holotype (Fig. 2A, C, E���F). The paratype and referred specimens have a dark inverted triangle pattern on the parietal-orbital-scapular region. The paratype has orange-reddish coloration on the throat, chest, and ventral surface of the fingers and toes (Fig 2C, D). The juvenile specimens (MZB Amph 32593 and UTA A-66177) have darker brown throats and chests. Comparisons. Megophrys selatanensis sp. nov. is morphologically most similar to M. montana than to other geographically related congeners. Megophrys selatanensis sp. nov. has elongated dorsolateral folds that extend from the parietoscapular region to the groin, type I, as M. montana, M. parallela, M. lancip, M. nasuta, M. kalimantanensis Munir, Hamidy, Matsui, Iskandar, Sidik, and Shimada, and M. stejnegeri Taylor. Megophrys selatanensis sp. nov. differs from M. montana by having a shorter rostral appendage���SAL 0.3���0.6% SVLh (vs. SAL reaching 3.5% SVLh: Munir et al., 2018), shorter palpebral projection���EHL 21.3���24.3% UEWh (vs. reaching 48.4% UEWh: Munir et al., 2018), shorter ratio of foot to thigh���FL 96.6���99.4% TL (vs. FL 106.2���115.6% TL: Munir et al., 2018), and toes webbed only at base (vs. more developed web on second to fifth toes: Munir et al., 2018). Megophrys selatanensis sp. nov. differs from M. parallela by having elongated dorsolateral folds reaching the groin (vs. only reaching two-thirds of trunk), females being large���known adult female 79.1 mm SVL (vs. SVL ��� 58.3 mm: Munir et al., 2018), a rostral appendage present (vs. absent: Inger and Iskandar, 2005; Munir et al., 2018), shorter ratio of foot to thigh���FL 96.6���99.4% TL (vs. FL 105.4���112.5% TL: Munir et al., 2018). The new species differs from M. lancip by having a less developed rostral appendage���SAL 0.3���0.6% SVLh (vs. SAL 1.4���4.1% SVLh: Munir et al., 2018), a shorter head length���HLh 38.0���38.5% SVLh (vs. HLh 42.1���44.6% SVLh: Munir et al., 2018), slightly wider head ratio to its length���HW 119.3���126.4% HLh (vs. HW 103.2���116.0% HLh: Munir et al., 2018), relatively shorter ratio of feet to thigh���FL 96.6���99.4%TL (vs. FL 102.5���109.8% TL: Munir et al., 2018), toes webbed only at base (vs. rather developed on third, fourth and fifth toes: Munir et al., 2018). ......continued on the next page Megophrys selatanensis sp. nov. differs from M. nasuta by the absence of additional lateral folds on flanks (vs. presence: Munir et al., 2018, 2019), shorter acute rostral appendage���SAL 0.3���0.6% SVLh (vs. acuminate rostral appendage, SAL 1.8���9.2% SVLh: Munir et al., 2019), shorter acute palpebral projection���EHL 2.2���3.4% SVLh (vs. acuminate palpebral projection, EHL 4.4���13.0% SVLh: Munir et al., 2019), shorter snout to nostril length���SNLh 3.7���5.1% SVLh (vs. SNLh 5.1���13.8% SVLh: Munir et al., 2019), shorter head length���HLh 38.0���40.5% SVLh (vs. HLh 39.5���51.7.% SVLh: Munir et al., 2019), smaller ratio of palpebral projection to upper eyelid width���EHL 21.3���25.7% UEWh (vs. EHL 32.7���61.4% UEWh: Munir et al., 2019), and relatively wider head���HW 119.3���126.4% HLh (vs. HW 92.9���119.5% HLh: Munir et al., 2019). From M. kalimantanensis, the new species differs by the absence of additional lateral folds on flanks (vs. presence: Munir et al., 2019), female being smaller���known adult female 79.1 mm SVL (vs. 109.1���116.4 mm SVL: Munir et al., 2019), adpressed tibiotarsal articulation reaching posterior corner of eye in female (vs. posterior of tympanum, in female: Munir et al., 2019). From M. stejnegeri Taylor, Megophrys selatanensis sp. nov. differs by having the dorsolateral folds extending to the groin (vs. maximum of two-thirds length of trunk: Inger, 1954), female being slightly larger���known adult female 79.1 mm SVL (vs. SVL 57.0��� 77.8 mm: Taylor, 1920; Inger 1954), a rostral appendage present (vs. absent: Taylor, 1920; Inger, 1954), vomerine teeth present (vs. absent: Taylor, 1920; Inger, 1954), tympanum vertically elongated (vs. slightly rounded, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016), and smooth skin on the temporal region (vs. tuberculate, see figure 37 by Diesmos et al., 2015 and figure 18 by Sanguila et al., 2016). Megophrys selatanensis sp. nov. is distinguished from M. kobayashii Malkmus and Matsui by having type I dorsolateral folds (vs. dorsolateral folds extended from parietoscapular region to mid-body, type II, see Table 3), additional lateral folds on flanks absent (vs. present: Malkmus and Matsui, 1997; Munir et al., 2018), female being smaller���known adult female 79.1 mm SVL (vs. SVL 99.0���109.0 mm: Malkmus and Matsui, 1997; Inger and Stuebing, 2005), a rostral appendage present (vs. absent: Malkmus and Matsui, 1997), and smaller and fewer tubercles on the body flanks (vs. larger and numerous: Malkmus and Matsui, 1997). From M. ligayae Taylor, the new species differs in having type I dorsolateral folds (vs. type II: Taylor, 1920; Inger, 1954; Munir et al., 2018), no additional lateral folds on flanks (vs. present: Taylor, 1920; Inger, 1954; Munir et al., 2018), female being smaller���known adult female 79.1 mm SVL (vs. SVL 90.0 mm: Inger, 1954), tibiotarsal articulation reaching posterior corner of eye (vs. tympanum: Taylor, 1920), and toes webbed only at base (vs. rather developed web on third, fourth, and fifth toes, see figure 37C Diesmos et al., 2015). From M. edwardinae Inger, the new species differs by having type I dorsolateral folds (vs. absent: Inger, 1989), a rostral appendage present (vs. absent: Inger, 1989), vomerine teeth present (vs. absent: Inger, 1989), and smooth dorsal skin surface with low and dense tubercles (vs. dorsal surface with scattered round and elongated tubercles: Inger, 1989). Megophrys selatanensis sp. nov. can be distinguished from M. baluensis Boulenger by having type I dorsolateral folds (vs. dorsolateral folds formed by a series of elongated tubercle, type III: Inger and Stuebing, 2005: Munir et al., 2018), female being slightly larger���known adult female 79.1 mm SVL (vs. SVL ��� 70 mm: Inger, 1966; Inger et al., 1995), a rostral appendage present (vs. absent: Boulenger, 1899; Inger, 1966; Inger et al., 1995), having a triangular palpebral projection (vs. small palpebral projection like tubercle: Inger, 1989; Inger et al., 1995), having smaller and fewer tubercles on body flanks (vs. larger and numerous: Inger et al., 2017), and adpressed tibiotarsal articulation reaching posterior of eye (vs. shoulder: Boulenger, 1899). Megophrys selatanensis sp. nov. differs from M. dringi Inger, Stuebing, and Tan, M. aceras Boulenger, and M. longipes Boulenger by the absence of Y, X,>���et al., 1995, see Table 3). Furthermore, from M. dringi the new species being larger in female���known adult female 79.1 mm SVL (vs. SVL 55 mm: Inger et al., 1995; Inger and Stuebing, 2005), having a stocky body (vs. slender: Inger et al., 1995), a rostral appendage present (vs. absent: Inger et al., 1995), having a triangular palpebral projection (vs. small like tubercle: Inger et al., 1995), vomerine teeth present (vs. absent: Inger et al., 1995), distinct tympanum (vs. partially obscured: Inger et al., 1995). The new species differs from M. aceras by having a stocky body (vs. slender: Boulenger, 1903), a rostral appendage present (vs. absent: Boulenger, 1903), tibiotarsal articulation reaching to posterior corner of eye (vs. shoulder, angle of jaws or temporal area: Taylor, 1962), and toes webbed only at base (vs. rather possessing developed web between third, fourth, and fifth toes: Taylor, 1962; see figure 5 in Munir et al., 2018). From M. longipes, the new species differs being larger in female���known adult female 79.1 mm SVL (vs. SVL 49.0���65.0 mm: Inger et al., 1995), having a stocky body (vs. slender: Boulenger, 1885), a rostral appendage present (vs. absent: Boulenger, 1885), having a triangular palpebral projection (vs. small like tubercle: Boulenger, 1885; Taylor, 1962), shorter thigh���TL 0.41���0.44 SVL (vs. TL 0.54���0.65 SVL: Inger et al., 1995; Inger and Iskandar, 2005) and tibiotarsal articulation reaching posterior corner of eye (vs. far beyond tip of snout: Boulenger, 1885; Taylor, 1962). Distribution and natural history. The holotype and one referred specimen (MZB Amph 32593) were collected in leaf litter on the edge of an old secondary forest in proximity to a coffee plantation and above a geothermal station in the province of Lampung, while the paratype (UTA A-66176) and the other referred specimen (UTA A- 66177) were collected from an old secondary forest in the green corridor between the Gunung Patah and Gunung Agung. These locations are about 170 km apart along the eastern slope of the southern Barisan range of Sumatra. The tadpoles (ENS 14306 and ENS 14737) were collected from a small rocky stream in the old secondary forest near Lake Ranau, Lampung, 1108���1487 m a.s.l. (description of the tadpole of this new species is pending as we are preparing another manuscript to present detailed description of Sumatran Megophrys tadpoles). Most of the samples of M. selatanensis sp. nov. were collected from a small patched forest area in the eastern Bukit Barisan Selatan Mountain ranges, which is threatened by high deforestation. The exact distribution, population, breeding behavior, call, and other ecological information are unknown. The following anuran species have been found sympatrically with the new species, at Ngarip, Lampung: Indosylvirana nicobariensis (Stoliczka); Chalcorana chalconota (Schlegel); Wijayarana sumatrana (Yang); Hylarana erythraea (Schlegel); Ingerophrynus biporcatus (Gravenhorst); Leptophryne borbonica (Tschudi); Limnonectes kuhlii (Tschudi); L. macrodon (Dum��ril and Bibron); Limnonectes sp.; Microhyla gadjahmadai Atmaja, Hamidy, Arisuryanti, Matsui, Smith; M. palmipes Boulenger; Pelophryne sp.; Philautus larutensis (Boulenger); P. polymorphus Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; P. thamyridion Wostl, Riyanto, Hamidy, Kurniawan, Smith, and Harvey; Phrynoides asper (Gravenhorst); Polypedates leucomystax (Gravenhorst); Rhacophorus achantharrhena, Published as part of Munir, Misbahul, Nishikawa, Kanto, Hamidy, Amir & Smith, Eric N., 2021, Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from Sumatra, Indonesia, pp. 503-529 in Zootaxa 5057 (4) on pages 507-514, DOI: 10.11646/zootaxa.5057.4.3, http://zenodo.org/record/5598859, {"references":["Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24 (1 - 4), 6 - 21.","Munir, M., Hamidy, A., Farajallah, A. & Smith, E. N. (2018) A new Megophrys Kuhl and van Hasselt (Amphibia: Megophryidae) from southwestern Sumatra, Indonesia. Zootaxa, 4442 (3), 389 - 412. https: // doi. org / 10.11646 / zootaxa. 4442.3.3","Inger, R. F. & Iskandar, D. T. (2005) A collection of amphibians from West Sumatra, with description of new species of Megophrys (Amphibia: Anura). The Raffles Bulletin of Zoology, 53 (1), 133 - 142.","Munir, M., Hamidy, A., Matsui, M., Iskandar, D. T., Sidik, I. & Shimada, T. (2019) A new species of Megophrys Kuhl & van Hasselt (Amphibia: Megophryidae) from Borneo allied to M. nasuta (Schlegel, 1858). Zootaxa, 4679 (1), 1 - 24. https: // doi. org / 10.11646 / zootaxa. 4679.1.1","Inger, R. F. (1954) The systematics and zoogeography of Philippine Amphibia. Fieldiana Zoology, 33 (4), 183 - 531. https: // doi. org / 10.5962 / bhl. title. 5571","Taylor, E. H. (1920) Philippine Amphibia. The Philippine Journal of Science, 16 (3), 213 - 359. https: // doi. org / 10.5962 / bhl. part. 4751","Diesmos, A. C., Watters, J. L., Huron, N. A., Davis, D. R., Alcala, A. C., Crombie R. I., Afuang, L. E., Gee-das, G., Sison, R. V., Sanguila, M. B., Penrod, M. L., Labonte, M. J., Davey C. S., Leone E. A., Diesmos M. L., Sy, M. Y., Welton, L. J., Brown R. M. & Siler C. D. (2015) Amphibians of the Philippines, Part I: Checklist of the Species. Proceedings of the California Academy of Science, 62 (20), 457 - 539.","Sanguila, M. B., Cobb, K. A, Siller C. D., Diesmos, A. C., Alcala A. C. & Brown R. M. (2016) The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands. ZooKeys, 624, 1 - 32. https: // doi. org / 10.3897 / zookeys. 624.9814","Malkmus, R. & Matsui, M. (1997) Megophrys kobayashii, ein neuer pelobatider frosch vom Mount Kinabalu, Sauria, 19, 31 - 37.","Inger, R. F. & Stuebing, R. B. (2005) A field guide to the frogs of Borneo. 2 nd Edition. Natural History Publication Borneo Sdn. Bhd. Kota Kinabalu, Sabah, 201 pp.","Inger, R. F. (1989) Four new species of frogs from Borneo. Malayan Nature Journal, 42, 229 - 243.","Inger, R. F. (1966) The systematic and zoogeography of the amphibians of Borneo. Fieldiana Zoology, 52, 1 - 402. https: // doi. org / 10.5962 / bhl. title. 3147","Inger, R. F., Stuebing, R. B. & Tan, F. L. (1995) New species and new records of anurans from Borneo. The Raffles Bulletin of Zoology, 43 (1), 115 - 131.","Boulenger, G. A. (1899) Description of three new reptiles and a new batrachian from Mount Kinabalu, North Borneo. Annals and Magazine of Natural History, 7 (4), 451 - 454. https: // doi. org / 10.1080 / 00222939908678228","Inger, R. F., Stuebing, R. B., Grafe, T. U. & Dehling, J. M. (2017). A field guide to the Frogs of Borneo. Third edition. Natural History Publication Borneo. Kinabalu, 236 pp.","Boulenger, G. A. (1885) Description of a new frog of the genus Megalophrys. Proceedings of the scientific meeting of the Zoological Society of London, 53 (4), 850 - 850. https: // doi. org / 10.1111 / j. 1096 - 3642.1885. tb 02927. x","Boulenger, G. A. (1903) Report on the batrachians and reptiles. In: Annandale, N. & Robinson, H. C. (Eds.), Fasciculi Malayenses. Zoology. Part I. Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States 1901 - 1902. Longmans, Green & Co, London, pp. 131 - 176.","Taylor, E. H. (1962) The amphibian fauna of Thailand. The University of Kansas Science Bulletin, 43 (8), 265 - 599. https: // doi. org / 10.5962 / bhl. part. 13347"]}

Das, Abhijit, Smith, Eric N., Sidik, Irvan, Sarker, Goutam C., Boruah, Bitupan, Patel, Naitik G., Deepak, V. (2021): Hidden in the plain sight: a new species of Rhabdophis (Serpentes: Natricinae) from the Rhabdophis himalayanus complex. Zootaxa 5020 (3): 401-433, DOI: https://doi.org/10.11646/zootaxa.5020.3.1

Sigalegalephrynus gayoluesensis sp. nov. Figs. 2 D���F, 4B, 5B, 6B Holotype. Museum Zoologicum Bogoriense of Amphibian Collection, MZB.Amph.30411 (field number ENS 19527). An adult male from above the Desa (Village) Kenyaran Pantan Cuaca, Kabupaten (Regency) Gayo Lues, Provinsi Aceh, Indonesia, 4.22588��N, 97.18915��E, 1850 m. a.s.l. (Fig. 3). Collected by Elijah Wostl, Ahmad Muammar Khadafi, and Syaripudin on 9 August 2015 at 21:20h. Paratypes (3). The University of Texas at Arlington Amphibian collection number UTA A-65490, Museum Zoologicum Bogoriense of Amphibian Collections, MZB.Amph.26035, adult males; MZB.Amph.26037, adult female. Collected from near to the collection locality of the holotype, 4.22580��N, 97.1886��1E, 1844 m. a.s.l. (Fig. 3). Collected by Elijah Wostl, Ahmad Muammar Khadafi, and Syaripudin on 9 August 2015 at 21:05h. Referred specimens (8). Collection locality very close to the types. UTA A-65488���489, 65789 (subadult and two juveniles, respectively, 1827 m. a.s.l., 4.2239��N, 97.18718��E); 65790 (subadult, 1826 m. a.s.l., 4.22487��N, 97.18769��E); and MZB.Amph. 26032 (juvenile, 1827 m. a.s.l., 4.22357��N, 97.186551�� E); 26033 (juvenile, 1827 m. a.s.l., 4.2239��N, 97.18718��E); 26034, 26036 (two juveniles, 1826 m. a.s.l., 4.22487��N, 97.18769��E). Etymology. The specific epithet refers to the Gayo Lues Highlands, where this new species was found. Suggested Common name. Gayo Lues Highland���s Puppet Toad; Indonesia name: Kodok-wayang gayolues Diagnosis. Sigalegalephrynus gayoluesensis sp. nov. can be identified from its congeners by a unique combination of characters: (1) medium-size (adult males 25.65���26.49 mm SVL); (2) lacking parotoid glands; (3) tympanum visible, with elevated annulus, and not encircled by sharply raised spinose tubercles; (4) naris closer to tip of snout than to eye; eye-naris distance 6.4.0% (7%) of SVL; naris-snout distance 1% (1.9%) of SVL; (5) fingertips truncated and expanded (except finger I); (6) tips of toe I, II and III are rounded; tips of toes IV and V truncated but not expanded; (7) rudimentary webbing in hands, moderate in feet; (8) adult male dorsal coloration dark brown, with prominent whitish diamond shaped suprascapular marking; (9) dorsum lacking medial dark band; (10) upper lip with prominent alternating dark brown and white marks; (11) flanks with stroke of dark brown (demarcated by thin white lines on top and bottom), extending from orbit to inguinal area; (12) dorsal surface lightly tuberculate, with round tubercles; (13) venter pinkish���white, with black maculation; (14) interocular distance 43% (44%) of head width; (15) nuptial pads dark brown, with black���tipped spicules; (16) finger IV tip extending beyond distal (terminal) phalangeal articulation of finger III (when adpressed); (17) inner metacarpal tubercle �� length to outer metacarpal tubercle. Description of holotype and variation of paratypes (in parenthesis). Body moderately robust; head longer than wide, HL/HW =1.14 (1.11, 1.07, 1.02); head length 33% (34%, 31%, 34%) of SVL; head width 29% (31%, 29%, 33%) of SVL; snout length 10% (10%, 10%, 11%) of SVL; canthus rostralis concave; loreal area without tubercles, concave; eye length 10% (12%, 10%, 9%) of SVL; pupil circular; snout truncate in dorsal view, protruding in lateral view, sloping back towards mouth; tympanum distinct and rounded, with annulus, but not surrounded by large tubercles; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval shaped and longer than wide; skin of dorsum finely shagreened, with few large and scattered tubercles; tubercles rounded, without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with anastomosis; circumcloacal region is golden yellow. Arms robust; forearm length 31% (33%, 25%, 26%) of SVL; hand length 30% (31%, 26%, 27%) of SVL; relative length of Finger���IMeasurements (in mm). Holotype followed by paratype in parenthesis: SVL 26.49 (25.65, 26.07, 27.36); HL 8.72 (8.84, 8.06, 9.20); HW 7.67 (7.98, 7.53, 9.0); SNL 2.75 (2.5, 2.55, 3.0); ICD 4.30 (4.50, 4.50, 4.56); IND 2.20 (1.80, 2.0, 2.0); END 1.7 (1.8, 1.55, 2.12); NSD 0.25 (0.5, 0.5, 0.7); IOD 3.0 (3.5, 3.5, 4.0); EL 2.70 (3.0, 2.55, 2.55); TML 1.50 (1.55, 1.6, 1.47); FAL 8.24 (8.40, 6.5, 6.5); HAL 7.89 (7.86, 6.67, 7.5); THL 11.83 (11.34, 11.53, 12.07); TBL 10.68 (10.95, 10.29, 11.23); TRL 6.63 (6.48, 5.50, 5.50); FTL 10.83 (10.86, 10.90, 10.80); OMCL 1.0 (1.0, 10, 1.0); OMCW 1.0 (1.0, 1.0 1.0, 1.0); IMCL 0.75 (0.75, 0.65, 0.60); IMCW 0.50 (0.50, 0.50); IMTL 1.0 (1.0, 1.0, 1.0); IMTW 1.0 (1.0, 1.0, 1.0); F1L 1.5 (1.5, 1.5, 1.5); F2L 2.30 (2.25, 2.35 2.45); F3L 4.0(3.5, 3.5, 3.2); F4L 3.5 (3.0, 2.6, 2.55); T1L 2.0 (1.5, 1.5, 1.6); T2L 2.5 (2.0, 2.0, 2.0); T3L 3.2 (3.0, 3.3, 2.5); T4L 5.5 (5.0, 5.0, 4.5); T5L 4.0 (3.5, 3.5, 3.3); F3PD 1.25 (1.2, 1.0, 1.3); F3PB 1.0 (1.0, 0.9, 1.0). Color of holotype in life. (Figs. 2D, 2E, 2F). Dorsum predominantly brown, with suprascapular dark brown diamond-shaped marking encircled by light brown; flanks with alternate wide dark brown and narrow white stripes; wide whitish light-brown spot below eye; lore dark brown, with small light brown spot adjacent to anterior of orbit; iris golden yellow, heavily reticulated; dorsum of limbs dark brown, with dark-brown crossbars; large white tubercles present at point of posterior mandibular articulation; abdominal surface pink, with dark brown maculation; throat pinkish, with no maculation; underside of limbs pink, with dark brown maculation; iris golden yellow, with black reticulations. Color of holotype in preservative. In alcohol, pinkish coloration turned grey and venter whitish grey, maculated with dark brown blotches. Advertisement call. The call of the male holotype was recorded in the field and before collection. Ambient temperature at the time of recording was 17.2 ��C. The call is composed of 179 highly modulated notes given 0.245 seconds apart, on average (range, 0.140 ���0.907 seconds, SD �� 0.148 seconds). On average, each note is 0.049 seconds (range, 0.24���0.93, SD �� 0.18 seconds) in length and is composed of one distinct pulse. The average fundamental and dominant frequencies of the vocalization are 2474.361 (range, 2368.652���2627.051 Hz, SD �� 85.86 Hz) Hz and 4948.722 Hz (range, 4737.305���5254.102 Hz, SD ��171.7309 Hz) respectively (Fig. 7). Comparisons. Sigalegalephrynus gayoluesensis sp. nov. is likely restricted to the mountains of the Gayo Lues Regency of Aceh, Sumatera, and does not exist in sympatry with any other congener. Sigalegalephrynus gayoluesensis sp. nov. can be easily distinguished from S. mandailinguensis, S. minangkabauensis and S. harveyi sp. nov. by its smooth tubercles on the body (vs. sharp-tipped warty tubercles) and a diamond shaped marking on the dorsum (vs. hourglass in S. mandailinguensis, S. minangkabauensis and S. harveyi sp. nov., no hourglass or diamond shape mark in S. burnitelongensis sp. nov.). Sigalegalephrynus gayoluesensis sp. nov. can also be distinguished from S. burnitelongensis sp. nov. by its black anastomotic maculated throat and abdomen (vs immaculate throat and abdomen). Acoustic data is limited for Sigalegalephrynus species, the call of the holotype of S. gayoluesensis sp. nov., differs from that of S. mandailinguensis in duration (46.448 s vs 17.27 s), total number of notes (179 vs 62), notes per second (4 vs 6���7), average note length (0.49 s vs 0.029 s), average pause length between notes (0.245 s vs 0.012 s), and dominant frequency (4948.722 Hz vs 3400 Hz) (Fig 5). Distribution and natural history. Sigalegalephrynus gayoluesensis sp. nov. is known only from rain forest flanking a stream adjacent to the Takengon-Blangkejeren road above the village Kenyaran Pantan Cuaca, in the Gayo Lues Regency of the province of Aceh, between 1787 and 1796 m a.s.l. (Fig. 3). Both the holotype and paratype were found calling on broad smooth leaves, at 1.6 m and 3.8 m above ground, respectively. The call of the holotype was recorded. The call sounded similar to that of S. mandailinguensis at the time of recording. Both the holotype and paratype weighed 1.27 g. Our smallest juvenile of this species (UTA A-65789) was less than 1 cm (SVL 8.0 mm) in SVL and weighed 0.05 g., Published as part of Sarker, Goutam C., Wostl, Elijah, Thammachoti, Panupong, Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2019, New species, diversity, systematics, and conservation assessment of the Puppet Toads of Sumatra (Anura: Bufonidae: Sigalegalephrynus), pp. 365-391 in Zootaxa 4679 (2) on pages 377-379, DOI: 10.11646/zootaxa.4679.2.9, http://zenodo.org/record/3772640

Sigalegalephrynus harveyi sp. nov. Figs. 2 M–O, 4E, 5E, 6E Holotype. Museum Zoologicum Bogoriense of Amphibian Collection, MZB.Amph.30412 (field number ENS 18377). An adult male from Gunung Dempo above the Desa (Village) Kampung Empat, Kabupaten (Regency) Pagar Alam, Provinsi Sumatera Selatan, Sumatra, Indonesia, 4.040980ºS, 103.1481ºE, 1826 m a.s.l. (in all cases, datum = WGS84) (Fig. 3). Collected by Michael B. Harvey, Farits Alhadi, and Panupong Thammachoti on 8 July 2015, at 21:35h. Paratype. The University of Texas at Arlington Amphibian Collection UTA A-65474, an adult male. Collected from near to the collection locality of the holotype, 4.03923ºS, and 103.1473ºE, 1878 m a.s.l (Fig 3). Collected by Michael B. Harvey, Panupong Thammachoti, and Gilang Pradana on 10 July 2015, at 20:55h. Etymology. The specific epithet is a patronym in honor of Michael B. Harvey, one of the collectors of this new species, a friend, an outstanding herpetologist, and the co-Principal Investigator of the National Science Foundation (NSF) project that has contributed this and a significant number of other papers on the herpetofauna of Sumatra. Suggested Common Name. Harvey’s Puppet Toad, in English; Kodok-wayang Harvey, in Indonesian. Diagnosis. Sigalegalephrynus harveyi sp. nov. can be identified from its congeners by a unique combination of characters: (1) medium-sized (adult males 26.36–28.09 mm SVL) Sigalegalephrynus; (2) lacking parotoid glands; (3) tympanum visible, with elevated annulus encircled with sharply raised spinose tubercles; (4) naris closer to tip of snout than to eye; eye-naris distance 8.0% (9.3%) of SVL; naris-snout distance 2.8 % (2.1%) of SVL; (5) fingertips truncated (except finger I), but not expanded; (6) tips of toes I, II, III and V rounded, toe IV tip truncated, but not expanded; (7) webbing rudimentary in hands, moderate in feet; (8) dorsal coloration in adult males light brown, with a prominent hourglass shaped marking; (9) dorsum, lacking medial dark band; (10) prominent alternate dark brown and white marks on upper lip; (11) flanks with dark brown strokes (demarcated by thin white lines on top and bottom), extending from orbit to inguinal area; (12) dorsal surface very lightly tuberculate, with white tipped spinose tubercles; (13) venter golden–yellow, without dark maculation; (14) interocular distance 48% (52%) of head width; (15) nuptial pads white, with white–tipped spicules; (16) finger IV tip touches distal phalangeal articulation of finger III (when adpressed); (17) inner metacarpal tubercle equal in length to outer metacarpal tubercle. Description of holotype and variation in paratype (in parenthesis). Body slender; head longer than wide, HL/HW 1.11 (1.14); head length 30% (33%) of SVL; head width 27.0% (29%) of SVL; snout length 13% (14%) of SVL; canthus rostralis concave; loreal area smooth and concave; eye length 10% (10%) of SVL; pupil circular; snout slightly sloping back, towards mouth; snout mucronate, with prominent median keel, protruding in lateral view; tympanum distinct, rounded, with moderately developed annulus; interorbital space flat; cranial crests absent; jaws toothless; tongue tip oval shaped and longer than wide; dorsal skin tuberculate and rough, with mostly small and white tipped tubercles, lacking black keratinization; tympanum with elevated and distinct annulus, circled by large tubercles; no dorsolateral, paravertebral, or occipital folds; throat golden yellow; venter pinkish and goldenyellow, areolate in texture; circumcloacal region brownish yellow. Arms lanky, with poorly developed axillary membranes; forearm length 28% (28%) of SVL; hand length 27% (28%) of SVL; relative length of fingers I Thigh length 41% (43%) of SVL; tibia length 38% (41%) of SVL; tarsal length 24% (23%) of SVL; foot length 42% (41%) of SVL; relative lengths of toes I Measurements (in mm). Holotype followed by paratype in parentheses: SVL 26.36 (28.09); HL 8.0 (9.41); HW 7.18 (8.25); SNL 3.4 (3.8); ICD 4.2 (4.5); IND 2.0 (2.60); END 2.6 (12.1); NSD 0.06 (0.75); IOD 3.7 (4.0); EL 2.55 (2.70); TML 1.3 (2.1); FAL 7.25 (7.95); HAL 7.09 (7.80); THL 10.72 (11.96); TBL 10.11 (11.45); TRL 6.20 (6.41); FTL 10.96 (11.62); OMCL 1.0 (1.0); OMCW 1.0 (1.0); IMCL 1.0 (1.0); IMCW 0.38 (0.50); IMTL 1 (1); IMTW 1 (1); F1L 1.5 (1.8); F2L 2.25 (2.30); F3L 3.65 (3.98); F4L 2.75 (3.40); T1L 1.0 (1.5); T2L 1.5 (2); T3L 3.0 (3.0); T4L 5.0 (5.5); T5L 3.5 (4.0).; F3PD 0.75 (1.0); F3PB 0.75 (1.0). Color of holotype in life. Adult male holotype (Figs. 2M, 2N, 2O): dorsum predominantly brown, with an hourglass marking with whitish brown halo; iris brownish-yellow; flanks with alternate wide dark-brown and thin white oblique stripes, extending from post-ocular to inguinal areas; a very dark brown triangular blotch below anterior half of eye, with thin posterior white border that extends posteriorly on subocular rim; loreal region brown; dorsum of limbs darker than body dorsum, humeral and femoral segments without crossbars, distal segments with crossbars; area of posterior mandibular articulation with a whitish-yellow spot; lower flanks, inguinal, and circumcloacal regions golden-yellow; underside of body and head yellowish, with heavily melanized chest; ventral limb surfaces brown-salmon color; finger and toe tips pale salmon color, not melanized; iris bronze with black reticulations. Color of holotype in preservative. Differing slightly from that in life, specimens have lost the golden yellow and pinkish coloration, which has turned grey. Comparisons. Sigalegalephrynus harveyi sp. nov. differs from all congeners by the combination of possessing truncated but not expanded fingertips (except finger I) (vs truncated and highly expanded in S. gayoluesensis sp. nov. and S. burnitelongensis sp. nov.; truncated and moderately expanded in S. mandailinguensis; round in S. minankabauensis), and white tipped tubercles on the body (vs black tipped in S. mandailinguensis and S. minangkabauensis). Additionally, Sigalegalephrynus harveyi sp. nov. has a prominent hourglass shaped marking on the dorsum (vs missing in adult males of S. burnitelongensis sp. nov.), white-spiculed nuptial pads in adult males (vs black or dark brown tipped in S. mandailinguensis, S. gayoluesensis sp. nov., S. burnitelongensis sp. nov., unknown in S. minangkabauensis), an indistinct white loreal spot (vs very distinct in S. mandailinguensis and S. minangkabauensis, absent in S. gayoluesensis sp. nov. and S. burnitelongensis sp. nov.), inner and outer metacarpal tubercles of equal size (vs inner metacarpal tubercle larger in S. mandailinguensis and S. minangkabauensis, and smaller in S. gayoluesensis sp. nov. and S. burnitelongensis sp. nov., with respect to outer metacarpal tubercle), and Finger IV tip (when adpressed) not touching the terminal (distal) phalangeal articulation of Finger III (vs touching in S. mandailinguensis S. minankabauensis, and going beyond the articulation in S. gayoluesensis sp. nov.) (Fig. 6E). Distribution and natural history. Sigalegalephrynus harveyi sp. nov. is only known from montane cloud-forest on the south-eastern slopes of Gunung Dempo, from 1826 and 1878 m a.s.l. (Fig. 3), and does not exist sympatrically with any other congener. The holotype was found calling on a leaf about 2 m above ground. Call was not recorded. The paratype was inactive on a leaf, 10 cm above ground. The holotype was not weighed, the paratype was 1.09 g.

Sigalegalephrynus burnitelongensis sp. nov. Figs. 2 A���C, 4A, 5A, 6A Holotype. Museum Zoologicum Bogoriense of Amphibian Collection, MZB.Amph.30413 (field number ENS 18884), an adult male. Collected from a stream of Gunung Burni Telong near Desa (Village) Rambune, Kecamatan (Subdistrict) Timang Gajah, Kabupaten (Regency) Bener Meriah, Province of Aceh, Indonesia. 4.76455��N, 96.80138��E, 1519 m a.s.l (Fig. 3). Collected by Goutam C. Sarker, Irvan Sidik, Syaripudin and Muhammad Ikhsan on 9 August 2015 at 00:30h. vergence and bold cells represent divergence between the southern and northern Groups). ......continued on the next page......continued on the next page Paratypes (2). The University of Texas at Arlington Amphibian collection numbers UTA A-65788 and UTA A-65492, adult males. Collected from near to the collection locality of the holotype, 4.76455��N, 96.80138��E, 1519 m a.s.l. (Fig. 3). Collected by Goutam C. Sarker, Irvan Sidik, Syaripudin and Muhammad Ikhsan on 8 August 2015 at 23:50h. Referred specimens (33). All juveniles, UTA A���65493���509 (17), and MZB.Amph.26016���031 (16), same col- lection information as the types. Etymology. The specific epithet is an adjective in Aceh language derived from Burni, meaning Mountain (Gunung in Indonesian) and Telong, meaning burning (Bakar in Indonesian), or in Sanskrit Borni T��Loŋ, meaning ���burning mountain���. This is the local name for the volcano that is the type-locality of this new species, and the Latin suffix ��� ensis, denoting place. Suggested Common Name. Burning Mountain Puppet Toad, in English; Kodok-wayang burnitelong, in Indonesian. Diagnosis. Sigalegalephrynus burnitelongensis sp. nov. can be diagnosed from its congeners by a unique combination of characters: (1) small-size (males 21.73���23.06 mm SVL); (2) lacking parotoid glands; (3) tympanum visible, with elevated annulus not encircled by sharply raised spinose tubercles; (4) naris closer to tip of snout than to eye; eye-naris distance 6.3% (8.3%, 6.9%) of SVL; naris-snout distance 1.1 % (1.2% 1%) of SVL, (5) fingertips truncated but not expanded (except finger I); (6) tips of toe I, II and III rounded, truncated but not expanded on toe IV and V; (7) rudimentary webbing in hands, moderate in feet; (8) dorsum brown without any marking; (9) medial dorsal dark band absent; (10) lacking alternate dark brown and white markings on upper lip, or not prominent; (11) flanks lacking stroke of different color; (12) dorsum lightly tuberculate, tubercles round; (13) venter pinkish���yellow, without maculation and uniformly tuberculate, (14) interocular distance 44% (43%) of head width; (15) nuptial pads dark brown, with black���tipped spicules; (16) finger IV tip not reaching distal phalangeal articulation of finger III (when adpressed); (17) inner metacarpal tubercle �� of outer metacarpal tubercle in length. Description of holotype and variation of paratypes (in parenthesis). Body moderately robust; head slightly longer than wide, HL/HW = 1.03 (1.10, 1.02); head length 32% (32%, 31%) of SVL; head width 31% (29%, 31%) of SVL; snout length 11% (10, 11%) of SVL; canthus rostralis concave; loreal area slightly tuberculate and concave; eye length 10% (9%, 10%) of SVL; pupil circular; snout truncate in dorsal view and protruding (slightly sloping back towards mouth) in lateral view; tympanum round with distinct annulus; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval shaped and longer than wide; skin of dorsal surfaces slightly rough to finely shagreen, with few large, scattered, round tubercles; most tubercles small, almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smoother, with very small and round tubercles; circumcloacal region golden yellow. Arms robust, with moderately developed axillary membrane; forearm length 27% (27%, 26%) of SVL; hand length 27% (24%, 26%) of SVL; relative length of fingers: I Measurements (in mm). Holotype followed by paratypes in parenthesis. Finger III of right hand of paratype deformed, finger measurements of this specimen taken on left hand. SVL 22.18 (21.73, 23.06); HL 7.06 (6.96, 7.20); HW 6.84 (6.31, 7.04); SNL 2.40 (2.20, 2.45); ICD 3.70 (3.50, 3.80); IND 1.89 (1.91, 1.93); END 1.4 (1.80, 1.6); NSD 0.25 (0.26, 0.23); IOD 3.00 (3.10, 3.00); EL 2.20 (2.00, 2.25); TML 1.4 (1.45, 1.2); FAL 6.04 (5.90, 6.10); HAL 6.00 (5.30, 6.00); THL 9.70 (9.28, 9.82); TBL 9.09 (8.97, 9.24); TRL 4.55 (4.50, 5.51); FTL 9.00 (7.98, 9.34); OMCL 1.00 (1.00, 1.00); OMCW 1.00 (1.00, 1.00); IMCL 0.55 (0.50, 0.50); IMCW 0.75 (0.75, 0.75); IMTL 1.5(1.00, 0.90); IMTW 1.0(0.70, 0.80); F1L 0.80 (1.00, 1.00); F2L 1.60 (2.00, 2.00); F3L 3.15 (3.50, 3.50); F4L 2.10 (2.50, 2.50); T1L 1.00 (1.00, 1.00); T2L (1.40, 1.40); T3L 1.80 (2.00, 2.00); T4L 5.00 (4.00, 4.60); T5L 3.50 (3.0, 3.00); F3PD 0.90 (0.80, 1.00); F3PB 0.80 (0.60, 0.75). Color of holotype in life. Adult male holotype (Figs. 2A, 2B, 2C): dorsum predominantly light brown, lacking distinct markings; flanks brown, lacking oblique stripes; infraorbital part of maxilla with light-brown marking; lore light brown, with small dark-brown spot between orbit and naris; dorsum of limbs brown, lacking distinctive crossbar markings; moderately large white tubercles at posterior mandibular articulation; abdominal surface pink, with many yellow blotches; gular region, clavicular, and ventral surface of limbs pink, without yellow blotches; tips of fingers and toes blackish, with golden yellow blotches; iris golden yellow, with heavy black reticulations. Color of holotype in preservative. Differing slightly from that in life, pinkish coloration turned grey, and venter has turned whitish grey. Comparisons. Sigalegalephrynus burnitelongensis sp. nov. is restricted to Gunung Burni Telong, a volcano in Bener Meriah regency, Sumatra. Sigalegalephrynus burnitelongensis sp. nov. can be easily distinguished from all other congeners (including S. gayoluesensis sp. nov. from Gayo Lues Regency) by the lack of crossbar markings on the dorsal surface of the limbs. It differs from S. mandailinguensis, S. minangkabauensis and S. harveyi sp. nov. by its truncate (vs. mucronate) shaped snout in dorsal profile, stocky limbs (vs. lanky) smooth tubercles (vs. warty with sharp tips), and lacking an hourglass mark on the dorsum (vs. hourglass present). Distribution and natural history. Sigalegalephrynus burnitelongensis sp. nov. is known only from forest patches associated to small streams and surrounded by coffee plantations, at Gunung Burni Telong, near the village of Rambune in the province of Aceh, from 1519 m a.s.l. (Fig. 3). The holotype and paratype were found sitting on small leaves of shrubs 20 cm above ground. The holotype weighed 0.76 g, and the paratype 0.69 g. The smallest juvenile collected (UTA A-65505) was 9.6 mm in SVL and 0.06 g in weight., Published as part of Sarker, Goutam C., Wostl, Elijah, Thammachoti, Panupong, Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2019, New species, diversity, systematics, and conservation assessment of the Puppet Toads of Sumatra (Anura: Bufonidae: Sigalegalephrynus), pp. 365-391 in Zootaxa 4679 (2) on pages 370-377, DOI: 10.11646/zootaxa.4679.2.9, http://zenodo.org/record/3772640

Sigalegalephrynus harveyi sp. nov. Figs. 2 M���O, 4E, 5E, 6E Holotype. Museum Zoologicum Bogoriense of Amphibian Collection, MZB.Amph.30412 (field number ENS 18377). An adult male from Gunung Dempo above the Desa (Village) Kampung Empat, Kabupaten (Regency) Pagar Alam, Provinsi Sumatera Selatan, Sumatra, Indonesia, 4.040980��S, 103.1481��E, 1826 m a.s.l. (in all cases, datum = WGS84) (Fig. 3). Collected by Michael B. Harvey, Farits Alhadi, and Panupong Thammachoti on 8 July 2015, at 21:35h. Paratype. The University of Texas at Arlington Amphibian Collection UTA A-65474, an adult male. Collected from near to the collection locality of the holotype, 4.03923��S, and 103.1473��E, 1878 m a.s.l (Fig 3). Collected by Michael B. Harvey, Panupong Thammachoti, and Gilang Pradana on 10 July 2015, at 20:55h. Etymology. The specific epithet is a patronym in honor of Michael B. Harvey, one of the collectors of this new species, a friend, an outstanding herpetologist, and the co-Principal Investigator of the National Science Foundation (NSF) project that has contributed this and a significant number of other papers on the herpetofauna of Sumatra. Suggested Common Name. Harvey���s Puppet Toad, in English; Kodok-wayang Harvey, in Indonesian. Diagnosis. Sigalegalephrynus harveyi sp. nov. can be identified from its congeners by a unique combination of characters: (1) medium-sized (adult males 26.36���28.09 mm SVL) Sigalegalephrynus; (2) lacking parotoid glands; (3) tympanum visible, with elevated annulus encircled with sharply raised spinose tubercles; (4) naris closer to tip of snout than to eye; eye-naris distance 8.0% (9.3%) of SVL; naris-snout distance 2.8 % (2.1%) of SVL; (5) fingertips truncated (except finger I), but not expanded; (6) tips of toes I, II, III and V rounded, toe IV tip truncated, but not expanded; (7) webbing rudimentary in hands, moderate in feet; (8) dorsal coloration in adult males light brown, with a prominent hourglass shaped marking; (9) dorsum, lacking medial dark band; (10) prominent alternate dark brown and white marks on upper lip; (11) flanks with dark brown strokes (demarcated by thin white lines on top and bottom), extending from orbit to inguinal area; (12) dorsal surface very lightly tuberculate, with white tipped spinose tubercles; (13) venter golden���yellow, without dark maculation; (14) interocular distance 48% (52%) of head width; (15) nuptial pads white, with white���tipped spicules; (16) finger IV tip touches distal phalangeal articulation of finger III (when adpressed); (17) inner metacarpal tubercle equal in length to outer metacarpal tubercle. Description of holotype and variation in paratype (in parenthesis). Body slender; head longer than wide, HL/HW 1.11 (1.14); head length 30% (33%) of SVL; head width 27.0% (29%) of SVL; snout length 13% (14%) of SVL; canthus rostralis concave; loreal area smooth and concave; eye length 10% (10%) of SVL; pupil circular; snout slightly sloping back, towards mouth; snout mucronate, with prominent median keel, protruding in lateral view; tympanum distinct, rounded, with moderately developed annulus; interorbital space flat; cranial crests absent; jaws toothless; tongue tip oval shaped and longer than wide; dorsal skin tuberculate and rough, with mostly small and white tipped tubercles, lacking black keratinization; tympanum with elevated and distinct annulus, circled by large tubercles; no dorsolateral, paravertebral, or occipital folds; throat golden yellow; venter pinkish and goldenyellow, areolate in texture; circumcloacal region brownish yellow. Arms lanky, with poorly developed axillary membranes; forearm length 28% (28%) of SVL; hand length 27% (28%) of SVL; relative length of fingers I Measurements (in mm). Holotype followed by paratype in parentheses: SVL 26.36 (28.09); HL 8.0 (9.41); HW 7.18 (8.25); SNL 3.4 (3.8); ICD 4.2 (4.5); IND 2.0 (2.60); END 2.6 (12.1); NSD 0.06 (0.75); IOD 3.7 (4.0); EL 2.55 (2.70); TML 1.3 (2.1); FAL 7.25 (7.95); HAL 7.09 (7.80); THL 10.72 (11.96); TBL 10.11 (11.45); TRL 6.20 (6.41); FTL 10.96 (11.62); OMCL 1.0 (1.0); OMCW 1.0 (1.0); IMCL 1.0 (1.0); IMCW 0.38 (0.50); IMTL 1 (1); IMTW 1 (1); F1L 1.5 (1.8); F2L 2.25 (2.30); F3L 3.65 (3.98); F4L 2.75 (3.40); T1L 1.0 (1.5); T2L 1.5 (2); T3L 3.0 (3.0); T4L 5.0 (5.5); T5L 3.5 (4.0).; F3PD 0.75 (1.0); F3PB 0.75 (1.0). Color of holotype in life. Adult male holotype (Figs. 2M, 2N, 2O): dorsum predominantly brown, with an hourglass marking with whitish brown halo; iris brownish-yellow; flanks with alternate wide dark-brown and thin white oblique stripes, extending from post-ocular to inguinal areas; a very dark brown triangular blotch below anterior half of eye, with thin posterior white border that extends posteriorly on subocular rim; loreal region brown; dorsum of limbs darker than body dorsum, humeral and femoral segments without crossbars, distal segments with crossbars; area of posterior mandibular articulation with a whitish-yellow spot; lower flanks, inguinal, and circumcloacal regions golden-yellow; underside of body and head yellowish, with heavily melanized chest; ventral limb surfaces brown-salmon color; finger and toe tips pale salmon color, not melanized; iris bronze with black reticulations. Color of holotype in preservative. Differing slightly from that in life, specimens have lost the golden yellow and pinkish coloration, which has turned grey. Comparisons. Sigalegalephrynus harveyi sp. nov. differs from all congeners by the combination of possessing truncated but not expanded fingertips (except finger I) (vs truncated and highly expanded in S. gayoluesensis sp. nov. and S. burnitelongensis sp. nov.; truncated and moderately expanded in S. mandailinguensis; round in S. minankabauensis), and white tipped tubercles on the body (vs black tipped in S. mandailinguensis and S. minangkabauensis). Additionally, Sigalegalephrynus harveyi sp. nov. has a prominent hourglass shaped marking on the dorsum (vs missing in adult males of S. burnitelongensis sp. nov.), white-spiculed nuptial pads in adult males (vs black or dark brown tipped in S. mandailinguensis, S. gayoluesensis sp. nov., S. burnitelongensis sp. nov., unknown in S. minangkabauensis), an indistinct white loreal spot (vs very distinct in S. mandailinguensis and S. minangkabauensis, absent in S. gayoluesensis sp. nov. and S. burnitelongensis sp. nov.), inner and outer metacarpal tubercles of equal size (vs inner metacarpal tubercle larger in S. mandailinguensis and S. minangkabauensis, and smaller in S. gayoluesensis sp. nov. and S. burnitelongensis sp. nov., with respect to outer metacarpal tubercle), and Finger IV tip (when adpressed) not touching the terminal (distal) phalangeal articulation of Finger III (vs touching in S. mandailinguensis S. minankabauensis, and going beyond the articulation in S. gayoluesensis sp. nov.) (Fig. 6E). Distribution and natural history. Sigalegalephrynus harveyi sp. nov. is only known from montane cloud-forest on the south-eastern slopes of Gunung Dempo, from 1826 and 1878 m a.s.l. (Fig. 3), and does not exist sympatrically with any other congener. The holotype was found calling on a leaf about 2 m above ground. Call was not recorded. The paratype was inactive on a leaf, 10 cm above ground. The holotype was not weighed, the paratype was 1.09 g., Published as part of Sarker, Goutam C., Wostl, Elijah, Thammachoti, Panupong, Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2019, New species, diversity, systematics, and conservation assessment of the Puppet Toads of Sumatra (Anura: Bufonidae: Sigalegalephrynus), pp. 365-391 in Zootaxa 4679 (2) on pages 379-382, DOI: 10.11646/zootaxa.4679.2.9, http://zenodo.org/record/3772640

Key to the species of Sigalegalephrynus 1 Adult males have a stout body with stocky limbs, and dorsum with a white diamond shaped mark or unmarked (Figs. 4 A���B); snout truncated in dorsal profile, and tympanic annulus well developed and covered with sharply raised tubercles (Figs. 5 A��� B)................................................................................................. 2 - Adult males and juveniles with a gracile body and lanky limbs, and dorsum with an hourglass shaped mark (Figs. 4 C���E); snout moderately mucronated in dorsal profile, and tympanic annulus not covered by sharply raised tubercles (Figs. 5 C���E)...... 3 2 Adult males> 24 mm in SVL, a white diamond shaped mark present on dorso-scapular region, and venter maculated in adult males (Fig. 4B); subarticular tubercle of finger I as wide as width of inner metacarpal tubercle, tip of finger IV extending beyond distal phalangeal articulation of finger III, when addpressed (Fig. 6B)............................................................................................................................... S. gayoluesensis - Adult males S. burnitelongensis 3 Venter in adult males maculated or blotched (Figs. 4C, 4E); webbing between toes I and II not complete (Figs. 6C, 6E); posterior mandibular articulation with a white spot on each side, and post-tympanic region with black and white large tubercles (Figs. 5C, 5E)............................................................................................. 4 - Venter in juveniles yellow with black blotches (Fig. 4D); webbing between toes I and II complete (Fig. 6D); posterior mandibular articulation without a white spot on each side, and post-tympanic region with only white large tubercles; fingertips rounded (Fig. 5D)........................................................................... S. minangkabauensis 4 Adult males> 30 mm in SVL, venter in adult males maculated and anastomotic, and tubercles on body with dark brown or black keratinized tips (Fig. 4C); nuptial pads in adult males with black-tipped spicules (Fig. 5C); finger tips truncated and expanded (Fig. 6C)............................................................................ S. mandailinguensis - Adult males S. harveyi, Published as part of Sarker, Goutam C., Wostl, Elijah, Thammachoti, Panupong, Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2019, New species, diversity, systematics, and conservation assessment of the Puppet Toads of Sumatra (Anura: Bufonidae: Sigalegalephrynus), pp. 365-391 in Zootaxa 4679 (2) on pages 382-383, DOI: 10.11646/zootaxa.4679.2.9, http://zenodo.org/record/3772640

Sigalegalephrynus gayoluesensis sp. nov. Figs. 2 D–F, 4B, 5B, 6B Holotype. Museum Zoologicum Bogoriense of Amphibian Collection, MZB.Amph.30411 (field number ENS 19527). An adult male from above the Desa (Village) Kenyaran Pantan Cuaca, Kabupaten (Regency) Gayo Lues, Provinsi Aceh, Indonesia, 4.22588ºN, 97.18915ºE, 1850 m. a.s.l. (Fig. 3). Collected by Elijah Wostl, Ahmad Muammar Khadafi, and Syaripudin on 9 August 2015 at 21:20h. Paratypes (3). The University of Texas at Arlington Amphibian collection number UTA A-65490, Museum Zoologicum Bogoriense of Amphibian Collections, MZB.Amph.26035, adult males; MZB.Amph.26037, adult female. Collected from near to the collection locality of the holotype, 4.22580ºN, 97.1886º1E, 1844 m. a.s.l. (Fig. 3). Collected by Elijah Wostl, Ahmad Muammar Khadafi, and Syaripudin on 9 August 2015 at 21:05h. Referred specimens (8). Collection locality very close to the types. UTA A-65488−489, 65789 (subadult and two juveniles, respectively, 1827 m. a.s.l., 4.2239ºN, 97.18718ºE); 65790 (subadult, 1826 m. a.s.l., 4.22487ºN, 97.18769ºE); and MZB.Amph. 26032 (juvenile, 1827 m. a.s.l., 4.22357ºN, 97.186551º E); 26033 (juvenile, 1827 m. a.s.l., 4.2239ºN, 97.18718ºE); 26034, 26036 (two juveniles, 1826 m. a.s.l., 4.22487ºN, 97.18769ºE). Etymology. The specific epithet refers to the Gayo Lues Highlands, where this new species was found. Suggested Common name. Gayo Lues Highland’s Puppet Toad; Indonesia name: Kodok-wayang gayolues Diagnosis. Sigalegalephrynus gayoluesensis sp. nov. can be identified from its congeners by a unique combination of characters: (1) medium-size (adult males 25.65–26.49 mm SVL); (2) lacking parotoid glands; (3) tympanum visible, with elevated annulus, and not encircled by sharply raised spinose tubercles; (4) naris closer to tip of snout than to eye; eye-naris distance 6.4.0% (7%) of SVL; naris-snout distance 1% (1.9%) of SVL; (5) fingertips truncated and expanded (except finger I); (6) tips of toe I, II and III are rounded; tips of toes IV and V truncated but not expanded; (7) rudimentary webbing in hands, moderate in feet; (8) adult male dorsal coloration dark brown, with prominent whitish diamond shaped suprascapular marking; (9) dorsum lacking medial dark band; (10) upper lip with prominent alternating dark brown and white marks; (11) flanks with stroke of dark brown (demarcated by thin white lines on top and bottom), extending from orbit to inguinal area; (12) dorsal surface lightly tuberculate, with round tubercles; (13) venter pinkish–white, with black maculation; (14) interocular distance 43% (44%) of head width; (15) nuptial pads dark brown, with black–tipped spicules; (16) finger IV tip extending beyond distal (terminal) phalangeal articulation of finger III (when adpressed); (17) inner metacarpal tubercle ¾ length to outer metacarpal tubercle. Description of holotype and variation of paratypes (in parenthesis). Body moderately robust; head longer than wide, HL/HW =1.14 (1.11, 1.07, 1.02); head length 33% (34%, 31%, 34%) of SVL; head width 29% (31%, 29%, 33%) of SVL; snout length 10% (10%, 10%, 11%) of SVL; canthus rostralis concave; loreal area without tubercles, concave; eye length 10% (12%, 10%, 9%) of SVL; pupil circular; snout truncate in dorsal view, protruding in lateral view, sloping back towards mouth; tympanum distinct and rounded, with annulus, but not surrounded by large tubercles; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval shaped and longer than wide; skin of dorsum finely shagreened, with few large and scattered tubercles; tubercles rounded, without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with anastomosis; circumcloacal region is golden yellow. Arms robust; forearm length 31% (33%, 25%, 26%) of SVL; hand length 30% (31%, 26%, 27%) of SVL; relative length of Finger—I Thigh length 45% (44%, 44%, 44%) of SVL; tibia length 40% (43%, 39%, 41%) of SVL; tarsal length 25% (25%, 21%, 20%) of SVL; foot length 41% (42%, 42%, 39%) of SVL; relative lengths of toes—I Measurements (in mm). Holotype followed by paratype in parenthesis: SVL 26.49 (25.65, 26.07, 27.36); HL 8.72 (8.84, 8.06, 9.20); HW 7.67 (7.98, 7.53, 9.0); SNL 2.75 (2.5, 2.55, 3.0); ICD 4.30 (4.50, 4.50, 4.56); IND 2.20 (1.80, 2.0, 2.0); END 1.7 (1.8, 1.55, 2.12); NSD 0.25 (0.5, 0.5, 0.7); IOD 3.0 (3.5, 3.5, 4.0); EL 2.70 (3.0, 2.55, 2.55); TML 1.50 (1.55, 1.6, 1.47); FAL 8.24 (8.40, 6.5, 6.5); HAL 7.89 (7.86, 6.67, 7.5); THL 11.83 (11.34, 11.53, 12.07); TBL 10.68 (10.95, 10.29, 11.23); TRL 6.63 (6.48, 5.50, 5.50); FTL 10.83 (10.86, 10.90, 10.80); OMCL 1.0 (1.0, 10, 1.0); OMCW 1.0 (1.0, 1.0 1.0, 1.0); IMCL 0.75 (0.75, 0.65, 0.60); IMCW 0.50 (0.50, 0.50); IMTL 1.0 (1.0, 1.0, 1.0); IMTW 1.0 (1.0, 1.0, 1.0); F1L 1.5 (1.5, 1.5, 1.5); F2L 2.30 (2.25, 2.35 2.45); F3L 4.0(3.5, 3.5, 3.2); F4L 3.5 (3.0, 2.6, 2.55); T1L 2.0 (1.5, 1.5, 1.6); T2L 2.5 (2.0, 2.0, 2.0); T3L 3.2 (3.0, 3.3, 2.5); T4L 5.5 (5.0, 5.0, 4.5); T5L 4.0 (3.5, 3.5, 3.3); F3PD 1.25 (1.2, 1.0, 1.3); F3PB 1.0 (1.0, 0.9, 1.0). Color of holotype in life. (Figs. 2D, 2E, 2F). Dorsum predominantly brown, with suprascapular dark brown diamond-shaped marking encircled by light brown; flanks with alternate wide dark brown and narrow white stripes; wide whitish light-brown spot below eye; lore dark brown, with small light brown spot adjacent to anterior of orbit; iris golden yellow, heavily reticulated; dorsum of limbs dark brown, with dark-brown crossbars; large white tubercles present at point of posterior mandibular articulation; abdominal surface pink, with dark brown maculation; throat pinkish, with no maculation; underside of limbs pink, with dark brown maculation; iris golden yellow, with black reticulations. Color of holotype in preservative. In alcohol, pinkish coloration turned grey and venter whitish grey, maculated with dark brown blotches. Advertisement call. The call of the male holotype was recorded in the field and before collection. Ambient temperature at the time of recording was 17.2 ºC. The call is composed of 179 highly modulated notes given 0.245 seconds apart, on average (range, 0.140 –0.907 seconds, SD ± 0.148 seconds). On average, each note is 0.049 seconds (range, 0.24–0.93, SD ± 0.18 seconds) in length and is composed of one distinct pulse. The average fundamental and dominant frequencies of the vocalization are 2474.361 (range, 2368.652–2627.051 Hz, SD ± 85.86 Hz) Hz and 4948.722 Hz (range, 4737.305–5254.102 Hz, SD ±171.7309 Hz) respectively (Fig. 7). Comparisons. Sigalegalephrynus gayoluesensis sp. nov. is likely restricted to the mountains of the Gayo Lues Regency of Aceh, Sumatera, and does not exist in sympatry with any other congener. Sigalegalephrynus gayoluesensis sp. nov. can be easily distinguished from S. mandailinguensis, S. minangkabauensis and S. harveyi sp. nov. by its smooth tubercles on the body (vs. sharp-tipped warty tubercles) and a diamond shaped marking on the dorsum (vs. hourglass in S. mandailinguensis, S. minangkabauensis and S. harveyi sp. nov., no hourglass or diamond shape mark in S. burnitelongensis sp. nov.). Sigalegalephrynus gayoluesensis sp. nov. can also be distinguished from S. burnitelongensis sp. nov. by its black anastomotic maculated throat and abdomen (vs immaculate throat and abdomen). Acoustic data is limited for Sigalegalephrynus species, the call of the holotype of S. gayoluesensis sp. nov., differs from that of S. mandailinguensis in duration (46.448 s vs 17.27 s), total number of notes (179 vs 62), notes per second (4 vs 6–7), average note length (0.49 s vs 0.029 s), average pause length between notes (0.245 s vs 0.012 s), and dominant frequency (4948.722 Hz vs 3400 Hz) (Fig 5). Distribution and natural history. Sigalegalephrynus gayoluesensis sp. nov. is known only from rain forest flanking a stream adjacent to the Takengon-Blangkejeren road above the village Kenyaran Pantan Cuaca, in the Gayo Lues Regency of the province of Aceh, between 1787 and 1796 m a.s.l. (Fig. 3). Both the holotype and paratype were found calling on broad smooth leaves, at 1.6 m and 3.8 m above ground, respectively. The call of the holotype was recorded. The call sounded similar to that of S. mandailinguensis at the time of recording. Both the holotype and paratype weighed 1.27 g. Our smallest juvenile of this species (UTA A-65789) was less than 1 cm (SVL 8.0 mm) in SVL and weighed 0.05 g.

Sigalegalephrynus burnitelongensis sp. nov. Figs. 2 A–C, 4A, 5A, 6A Holotype. Museum Zoologicum Bogoriense of Amphibian Collection, MZB.Amph.30413 (field number ENS 18884), an adult male. Collected from a stream of Gunung Burni Telong near Desa (Village) Rambune, Kecamatan (Subdistrict) Timang Gajah, Kabupaten (Regency) Bener Meriah, Province of Aceh, Indonesia. 4.76455ºN, 96.80138ºE, 1519 m a.s.l (Fig. 3). Collected by Goutam C. Sarker, Irvan Sidik, Syaripudin and Muhammad Ikhsan on 9 August 2015 at 00:30h. vergence and bold cells represent divergence between the southern and northern Groups). ......continued on the next page......continued on the next page Paratypes (2). The University of Texas at Arlington Amphibian collection numbers UTA A-65788 and UTA A-65492, adult males. Collected from near to the collection locality of the holotype, 4.76455ºN, 96.80138ºE, 1519 m a.s.l. (Fig. 3). Collected by Goutam C. Sarker, Irvan Sidik, Syaripudin and Muhammad Ikhsan on 8 August 2015 at 23:50h. Referred specimens (33). All juveniles, UTA A−65493−509 (17), and MZB.Amph.26016−031 (16), same col- lection information as the types. Etymology. The specific epithet is an adjective in Aceh language derived from Burni, meaning Mountain (Gunung in Indonesian) and Telong, meaning burning (Bakar in Indonesian), or in Sanskrit Borni TƏLoŋ, meaning “burning mountain”. This is the local name for the volcano that is the type-locality of this new species, and the Latin suffix – ensis, denoting place. Suggested Common Name. Burning Mountain Puppet Toad, in English; Kodok-wayang burnitelong, in Indonesian. Diagnosis. Sigalegalephrynus burnitelongensis sp. nov. can be diagnosed from its congeners by a unique combination of characters: (1) small-size (males 21.73–23.06 mm SVL); (2) lacking parotoid glands; (3) tympanum visible, with elevated annulus not encircled by sharply raised spinose tubercles; (4) naris closer to tip of snout than to eye; eye-naris distance 6.3% (8.3%, 6.9%) of SVL; naris-snout distance 1.1 % (1.2% 1%) of SVL, (5) fingertips truncated but not expanded (except finger I); (6) tips of toe I, II and III rounded, truncated but not expanded on toe IV and V; (7) rudimentary webbing in hands, moderate in feet; (8) dorsum brown without any marking; (9) medial dorsal dark band absent; (10) lacking alternate dark brown and white markings on upper lip, or not prominent; (11) flanks lacking stroke of different color; (12) dorsum lightly tuberculate, tubercles round; (13) venter pinkish–yellow, without maculation and uniformly tuberculate, (14) interocular distance 44% (43%) of head width; (15) nuptial pads dark brown, with black–tipped spicules; (16) finger IV tip not reaching distal phalangeal articulation of finger III (when adpressed); (17) inner metacarpal tubercle ¾ of outer metacarpal tubercle in length. Description of holotype and variation of paratypes (in parenthesis). Body moderately robust; head slightly longer than wide, HL/HW = 1.03 (1.10, 1.02); head length 32% (32%, 31%) of SVL; head width 31% (29%, 31%) of SVL; snout length 11% (10, 11%) of SVL; canthus rostralis concave; loreal area slightly tuberculate and concave; eye length 10% (9%, 10%) of SVL; pupil circular; snout truncate in dorsal view and protruding (slightly sloping back towards mouth) in lateral view; tympanum round with distinct annulus; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval shaped and longer than wide; skin of dorsal surfaces slightly rough to finely shagreen, with few large, scattered, round tubercles; most tubercles small, almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smoother, with very small and round tubercles; circumcloacal region golden yellow. Arms robust, with moderately developed axillary membrane; forearm length 27% (27%, 26%) of SVL; hand length 27% (24%, 26%) of SVL; relative length of fingers: I Thigh length 44% (43%, 43%) of SVL; tibia length 41% (41%, 40%) of SVL; tarsal length 21% (21%, 24%) of SVL; foot length 41% (37%, 41%) of SVL; relative lengths of toes—I Measurements (in mm). Holotype followed by paratypes in parenthesis. Finger III of right hand of paratype deformed, finger measurements of this specimen taken on left hand. SVL 22.18 (21.73, 23.06); HL 7.06 (6.96, 7.20); HW 6.84 (6.31, 7.04); SNL 2.40 (2.20, 2.45); ICD 3.70 (3.50, 3.80); IND 1.89 (1.91, 1.93); END 1.4 (1.80, 1.6); NSD 0.25 (0.26, 0.23); IOD 3.00 (3.10, 3.00); EL 2.20 (2.00, 2.25); TML 1.4 (1.45, 1.2); FAL 6.04 (5.90, 6.10); HAL 6.00 (5.30, 6.00); THL 9.70 (9.28, 9.82); TBL 9.09 (8.97, 9.24); TRL 4.55 (4.50, 5.51); FTL 9.00 (7.98, 9.34); OMCL 1.00 (1.00, 1.00); OMCW 1.00 (1.00, 1.00); IMCL 0.55 (0.50, 0.50); IMCW 0.75 (0.75, 0.75); IMTL 1.5(1.00, 0.90); IMTW 1.0(0.70, 0.80); F1L 0.80 (1.00, 1.00); F2L 1.60 (2.00, 2.00); F3L 3.15 (3.50, 3.50); F4L 2.10 (2.50, 2.50); T1L 1.00 (1.00, 1.00); T2L (1.40, 1.40); T3L 1.80 (2.00, 2.00); T4L 5.00 (4.00, 4.60); T5L 3.50 (3.0, 3.00); F3PD 0.90 (0.80, 1.00); F3PB 0.80 (0.60, 0.75). Color of holotype in life. Adult male holotype (Figs. 2A, 2B, 2C): dorsum predominantly light brown, lacking distinct markings; flanks brown, lacking oblique stripes; infraorbital part of maxilla with light-brown marking; lore light brown, with small dark-brown spot between orbit and naris; dorsum of limbs brown, lacking distinctive crossbar markings; moderately large white tubercles at posterior mandibular articulation; abdominal surface pink, with many yellow blotches; gular region, clavicular, and ventral surface of limbs pink, without yellow blotches; tips of fingers and toes blackish, with golden yellow blotches; iris golden yellow, with heavy black reticulations. Color of holotype in preservative. Differing slightly from that in life, pinkish coloration turned grey, and venter has turned whitish grey. Comparisons. Sigalegalephrynus burnitelongensis sp. nov. is restricted to Gunung Burni Telong, a volcano in Bener Meriah regency, Sumatra. Sigalegalephrynus burnitelongensis sp. nov. can be easily distinguished from all other congeners (including S. gayoluesensis sp. nov. from Gayo Lues Regency) by the lack of crossbar markings on the dorsal surface of the limbs. It differs from S. mandailinguensis, S. minangkabauensis and S. harveyi sp. nov. by its truncate (vs. mucronate) shaped snout in dorsal profile, stocky limbs (vs. lanky) smooth tubercles (vs. warty with sharp tips), and lacking an hourglass mark on the dorsum (vs. hourglass present). Distribution and natural history. Sigalegalephrynus burnitelongensis sp. nov. is known only from forest patches associated to small streams and surrounded by coffee plantations, at Gunung Burni Telong, near the village of Rambune in the province of Aceh, from 1519 m a.s.l. (Fig. 3). The holotype and paratype were found sitting on small leaves of shrubs 20 cm above ground. The holotype weighed 0.76 g, and the paratype 0.69 g. The smallest juvenile collected (UTA A-65505) was 9.6 mm in SVL and 0.06 g in weight.

Megophrys lancip sp. nov. (Figs. 3���5) Holotype. MZB Amph 22233 (field number ENS 14509) an adult male from the north-east of Desa Ngarip, Kecamatan Ulubelu, Kabupaten Tanggamus, Province of Lampung, Sumatra (5.31359��S, 104.53151��E, elevation 1010 m. a.s.l.), collected by Eric Nelson Smith, Amir Hamidy and Elijah Wostl, at 2005 h on 10 June 2013 (Fig. 3, 4A���E). Paratypes. UTA A-64763 (field number ENS 7579; formerly MZB Amph 12564) adult male from the Bengkulu to Kepahiang road, Kecamatan Taba penanjung, Kabupaten Bengkulu Tengah, Province of Bengkulu, Sumatra (3.68561��S, 102.53861��E, 620 m. a.s.l.), collected by Eric N. Smith and Michael B. Harvey, at 1740 h on 30 May 1996 (Fig. 5B); MZB Amph 3469, 14602, and 26738 (Fig. 4F���J), three adult females from Bukit Barisan Selatan National Park, at Desa Kubu Perahu, near Kota Liwa, Kecamatan Balik Bukit, Kabupaten Lampung Barat, Province of Lampung, Sumatra, collected by Andiek Fajar on 19 February 1998 (5.07870��S, 104.01189��E, 434 m. a.s.l.), by Inggar Ul-Hasanah on 17 September 2005 (5.07526��S, 104.04081��E, 550 m. a.s.l.), and by Anton Nurcahyo in 2005 (5.07526��S, 104.04081��E, 550 m. a.s.l.), respectively; MZB Amph 3 472 adult female from Bukit Barisan Selatan National Park, at Desa Sukaraja, Kecamatan Semaka, Kabupaten Tanggamus, Province of Lampung, Sumatra (5.51955��S, 104.45303��E, 526 m. a.s.l.), collected by Andiek Fajar on 6 November 1997. Etymology. The species name lancip is the Indonesian word for ���pointed���, used as an adjective, and reflects the extremely pointed rostral appendage of the new species. Suggested English common name. Pointed Horned Frog. Suggested Indonesian common name. Katak-tanduk lancip. Diagnosis. The new species is assigned to the genus and subgenus Megophrys based on our molecular analysis (Fig. 2) and the following morphological features: (1) absence of a distinct glandular fold forming a Y, X, or H on the parietoscapular region, possession of (2) a broad and depressed head, (3) pectoral and femoral glands, (4) eyelid appendages, (5) a vertical pupil, (6) a pair of conical spine at the corner of the mouth, and (7) a protruding snout (Delorme et al. 2006). Megophrys lancip sp. nov. is a medium-sized Megophrys (adult males SVLh 37.9���47.7 mm, n = 2; adult females SVLh 38.7���82.5 mm, n = 4) with the head shorter than the body (adult males HL 15.3���19.9 mm: 40.3���41.7% of SVLh, n = 2; adult females HL 16.0��� 35.4 mm: 41.4���42.9% of SVLh, n = 4), and broader compared to its length (adult males HL 87.7���95.8% of HW, n = 2; adult females 80.8���85.7% of HW, n = 4). The snout is moderate length (adult males SL 30.5���35.8% of HL, n = 2; adult females SL 29.4���33.6% of HL, n = 4), extremely pointed in dorsal view, protruding in lateral view, and with a sharp and pointed (acuminate) rostral appendage that projects beyond the lower jaw. The rostral appendage is moderate (adult males SNLh 3.0��� 3.3 mm, 7.0���8.0% to SVLh, n = 2; adult females SNLh 2.5���5.4 mm, 4.9���6.7% of SVLh, n = 4; adult males SAL; 1.2���1.4 mm, 2.9���3.1% of SVLh, n = 2; adult females SAL 0.6���3.44 mm, 1.5���4.1% of SVLh, n = 4); the upper eyelid has a triangular, short, and narrow-based appendage; the tympanum is distinct, small, oval, andoriented vertically; vomerine teeth are present; and the subarticular tubercle and outer metatarsal tubercles are absent. Description of Holotype (measurements in mm). Adult male; body stocky, robust (SVL 36.8; SVLh 37.9); head broad and depressed (HW 17.4: 46.0% of SVLh), slightly broader than long (HL 15.3: 40.3% of SVLh); snout extremely pointed from dorsal view, protruding in lateral view, tip with an acuminate sharp rostral appendage projection (SAL 1.2: 3.1% of SVLh), projecting beyond lower jaw; eye large and lateral on head, smaller (ED 5.0: 13.1% of SVLh) than snout length (SL 5.5: 14.4% of SVLh), eye diameter about 3.5 times of tympanum diameter (ED 5.0: 359.4% of TDH); canthus rostralis sharp, angular, concave; nostril below canthus on lateral view, distinctly closer to eye (NEL 1.9: 4.9% of SVLh) than to snout (SNL 2.3: 6.0% of SVLh); internarial distance (IND 3.4: 8.9% of SVLh) much shorter than interorbital distance (IOD 5.1: 13.5% of SVLh); upper eyelid with low tubercles and short narrow-base eyelid appendage projection (EHL 1.2: 2.9% of SVLh) much shorter than total width (UEWh 4.8: 12.7% of SVLh); a conical spine present at corner of mouth; tympanum distinct, oval, oriented vertically (TDV 2.2: 5.7% of SVLh; TDH 1.4: 3.6% of SVLh); vomerine teeth present; tongue not notched, without papilla. Forelimbs slender and short (FLL 25.5: 67.3% of SVLh), about half of hindlimbs (FLL 25.5: 49.0% of HLL); fingers moderately slender, unwebbed, without lateral fringes, with rounded and swollen tips; first finger (fin1L 3.1: 8.1% of SVLh) slightly longer than second (fin2L 3.0: 7.8% of SVLh) and fourth (fin4L 2.8: 7.5% of SVLh), third much longer (fin3L 5.0: 13.1% of SVLh); finger length formula IVSkin. Dorsal skin is smooth with low, sharply angular, tubercles from the snout to the vent, especially around the waist; the tubercles are denser laterally on the body and on the posterior surface of the thigh; upper eyelids with low conical tubercles; supratympanic fold distinct, angular, widening anteriorly, narrow posteriorly, extending from posterior of eye, curving down around upper border of tympanum and terminating above axilla; Y, X or H fold absent between parietoscapular area to level of axilla; dorsolateral fold extends from between eye and tympanum to the groin, wider from shoulder to groin; forelimb and hindlimb surface tuberculate and with transverse folds in forearm, tibia, and femur; a weakly developed tubercles ridge on outer of edge of forearm and tarsus forming a weak serration; ventral skin slightly tuberculate; skin on throat wrinkled, chest slightly tuberculate; ventral surface of limbs and abdomen smooth to weakly granular; a pair of white pectoral glands, small, conical and slightly raised; a pair of white femoral glands, moderately large, rounded and slightly raised, at mid-femur. Male with a pair of vocal slits on the posterior corners of the mouth, the single, external, vocal sac is indistinct on the preserved specimen; dark nuptial pads are present on the dorsomedial surface of the first and second fingers, from the base of the finger to the first terminal joint. Colour. In life (Fig. 3A���D), the entire dorsal surface of the head, body, forelimbs and hindlimbs is light brown; a darker brown inverse triangular marking bounded by a thin fold is present from the interorbital region to the posterior area of the head. The dorsal surface of the body, forelimbs and hindlimbs have indistinct, darker brown blotches. The dorsal surface of the lower arms, tibia and femur (hindlimbs) have dark brown transverse folds. The lateral side of the head, including the canthal area, is brown, darker around the nostrils and just posterior and below the canthus, lighter anterior and posterior to the eye, with dark markings above and below the skin bordering the eye; supratympanic fold light brown, with indistinct dark brown blotches. The entire ventral surface is light brown with a heavy dark transverse marking and dark blotches over the throat and on the chest and abdomen. Ventral surface of throat to chest is dark brown, while the abdomen, forelimbs and hindlimbs are light cream, with some small blotches. In preservative (Fig. 4A���E) aspects of the colour pattern remain, but the dorsum darkens. Variation (measurements in mm). Individuals of the type series are morphometrically generally similar (Table 4 and 5). Based on the individual specimens collected, females appear to be slightly larger and stockier than males (male SVL 36.8���46.3 mm [n = 2] vs female SVL 38.2���79.1 mm [n = 4]). When the hindlimb is bent forward along the body, the tibiotarsal joint usually reaches to the posterior corner of the eye in adult males and reaches to the posterior of tympanum in females. Only one female (MZB Amph 3 472) had the tibiotarsal joint reach the posterior of the eye. Individuals of the type series are only slightly variable in colouration. In preservative, the dorsal colouration in both males and females is relatively uniform light brown, but is completely dark brown in the single male paratype (MZB Amph 12564). The colouration of the ventral surface in both sexes is uniformly brown with dark blotches and dark longitudinal bands on the throat and chest. This longitudinal pattern is faded in one male (MZB Amph 12564). Comparisons. Megophrys lancip sp. nov. differs from members of subgenera Xenophrys, Atympanophrys, Panophrys, and M. dringi by the absence of a folded gland forming a Y, X or H directed backwards at the parietoscapular region (G��nther, 1864; Delorme et al. 2006) (versus presence) (Figs. 5A, 6C���D, Table 6). Megophrys lancip sp. nov. is distinguished from all known members of Sundaland and Philippine Megophrys (except for M. montana and M. nasuta), by the presence of a well -developed rostral appendage on the tip of the snout (versus absence in M. aceras, M. baluensis, M. dringi, M. edwardinae, M. longipes, M. kobayashii, and M. parallela) (Table 6, Fig. 5B���D). Megophrys lancip sp. nov. differs from Philippine M. stejnegeri by having a triangular-shaped eyelid appendage (versus a tubercle) and from M. ligayae by having a single dorsolateral fold that extends from the parietoscapular region to the groin (Table 6, type I) (versus a dorsolateral folds that extends from the parietoscapular region to the middle of the body [Table 6, type II]). Megophrys lancip sp. nov. can be distinguished from M. nasuta by absence of additional lateral fold that extends from the area behind the supratympanic fold to the groin (versus presence in M. nasuta) (Table 6; Fig. 5D), shorter rostral appendage SAL 1.5���4.1% of SVLh (versus SAL 1.2���9.2% of SVLh), and shorter eyelid appendage EHL 0.8���2.2 mm; 22.4% of UEWh (versus EHL 3.5���11.9 mm; 50.6% of UEWh) (Table 5; Fig. 5B, D) Megophrys lancip sp. nov. differs from M. montana by having a smaller body size, SVL 36.8���46.3 mm in adult males and 38.2���79.1 mm in adult females (versus 38.1���53.9 mm in adult males and 45.7���99.5 mm in adult females) (Table 4), longer snout appendage, SAL 1.5���4.1% of SVLh, (versus SAL 0.0���3.5% of SVLh), shorter eyelid appendage EHL 0.8���2.2 mm; 22.4% of UEWh (versus EHL 0.9���3.9 mm; 30.0% of UEWh), shorter hand length, 8.6���10.4 mm in adult males and 9.8���19.0 mm in adult females, (versus 10.4���14.4 mm in adult males and 11.0��� 23.6 mm in adult females), shorter hindlimb, HLL 52.0��� 58.8 mm in adult males and 43.4���98.3 mm in adult females, (versus 56.2���76.6 mm in adult males and 56.0��� 133.4 mm in adult females), tibio-tarsal articulation reached behind the eye in adult males and mostly reached behind the tympanum in adult females, (versus reached to the front of eye in adult males and reached the middle of eye in adult females), and less developed toe webbing I (1), II (1���1), III (2���2), IV (3���3), V (1 1/2) [versus I (1), II (1���1/2), III (1 3/4���1 1/2), IV (2 1/2���2 1/2), V(1)] (Table 6). TABLE ��. Variation in size anđ bođy proportions of male anđ female Megophrys from the Sunđalanđ anđ Philippines, mođifieđ from Taylor (1920), Inger (1954), Inger et al. (1995), Malkmus & Matsui (1997), anđ Inger & Iskanđar (2005). * for M. parallela, the measurement đoes not incluđe the rostral appenđage. ������continued on the next page ������continued on the next page ������continued on the next page Megophrys lancip sp nov. Megophrys montana Species RSAL RHLh RHW RAG REHL RLAL RHLL RFML RTL REHL/UEWh RTDH/ED RFL/TL RHW/HLh Species male (n = 28) female (n = 21) juvenile (n = 12) male (n =6) female (n = 2) RSAL 4.2 (1.5���8.7) 4.2 (1.8���9.2) 2.4 (1.2���5.4) ��� ��� ��� ��� RHLh 44.0 (39.5���49.7) 46.0 (40.6���52.8) 42.0 (36.9���46.7) 42.6 (39.1���44.3) 41.8 (41.6���42.0) RHW 46.4 (44.0���49.5) 48.4 (44.8���54.8) 45.2 (37.0���49.4) 46.6 (44.9���51.9) 48.5 (48.5���48.6) RAG 42.0 (33.5���47.0) 46.5 (39.6���57.2) 48.4 (42.7���51.6) 44.5 (40.1���47.3) 46.3 (40.1���52.6) REHL 8.3 (6.3���11.4) 8.9 (4.4���13.0) 9.6 (6.0���12.7) 2.8 (1.8���3.4) 3.7 (3.2���4.2) RLAL 46.6 (20.8���49.8) 32.4 (19.9���48.2) 23.3 (20.7���25.2) 49.3 (46.3���50.5) 47.8 (47.2���48.5) RHLL 121.0 (108.8���147.9) 116.8 (106.6���126.2) 119.0 (111.6���129.0) 148.4 (139.9���161.5) 147.2 (143.0���151.4) RFML 40.5 (36.0���48.1) 38.9 (33.2���41.9) 40.4 (37.5���44.8) 48.1 (44.3���51.0) 47.5 (45.2���49.7) RTL 36.0 (33.3���44.1) 35.1 (31.8���37.9) 37.1 (34.4���40.4) 43.9 (41.3���48.4) 44.4 (42.4���46.4) REHL /UEWh 50 (38.5���61.4) 53.6 (32.7���61.3) 48.4 (32.1���62.7) 21.9 (14.7���29.0) 30.6 (24.9���36.3) RTDH /ED 32.7 (22.2���43.9) 35.7 (16.3���52.8) 28.2 (10.9���34.5) 32.4 (22.1���35.8) 31.5 (29.6���33.3) RFL/TL 111.8 (103.0���118.2) 111.1 (98.8���121.3) 108.9 (101���113.3) 107.1 (105.4���112.5) 106.9 (106.6���107.2) RHW /HLh 106.4 (92.9���116.1) 103.9 (92.4���119.5) 106.8 (91.6���111.2) 111.7 (106.9���118.1) 116.1 (115.4���116.9) Megophrys parallela Relative values (R) of each character compared to SVL (* used HL, and HW compared to HL). * Fold forming Y, X or H on the parietoscapular region to the level of axilla. ** Dorsolateral fold shape: Type I, dorsolateral folds are elongated and extend from the parietoscapular region to the groin. Type II, dorsolateral folds extend from the parietoscapular region to the middle of the body Type III, multiple dorsolateral folds - at least three or four - and they are discontinuous, formed by a series of elongated tubercles. Type IV, dorsolateral folds are elongated and curve from the axillary region towards (and reaching) the posteriodorsal margin of tympanum. Distribution and Natural History. The new species is known from the provinces of Lampung and Bengkulu in southwestern Sumatra (Fig. 1). Larval, acoustic and other ecological data are unknown. The holotype was collected from a coffee plantation near the edge of secondary forest at 1010 m a.s.l. The male paratype from Bengkulu was collected from inside a rotten log on the forest floor at 620 m a.s.l. The female paratypes, collected from Bukit Barisan Selatan National Park, were collected from primary forest between 434 and 526 m a.s.l. This new species of Megophrys can be found sympatrically with M. nasuta in Kubu Prahu, Bukit Barisan Selatan National Park. Habitat loss and exploitation for the pet trade are likely be the main threats for the new species., Published as part of Munir, Misbahul, Hamidy, Amir, Farajallah, Achmad & Smith, Eric N., 2018, A New Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from southwestern Sumatra, Indonesia, pp. 389-412 in Zootaxa 4442 (3) on pages 395-408, DOI: 10.11646/zootaxa.4442.3.3, http://zenodo.org/record/1303723, {"references":["Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anoures). Alytes, 24 (1 - 4), 6 - 21.","Gunther, A. C. L. G. (1864) The reptiles of British India. The Ray Society, London, 452 pp. https: // doi. org / 10.5962 / bhl. title. 5012","Taylor, E. H. (1920) Philippine Amphibia. The Philippine Journal of Science, 16 (3), 213 - 359. https: // doi. org / 10.5962 / bhl. part. 4751","Inger, R. F. (1954) The systematics and zoogeography of Philippine Amphibia. Fieldiana Zoology, 33 (4), 183 - 531. https: // doi. org / 10.5962 / bhl. title. 5571","Inger, R. F., Stuebing, R. B. & Tan, F. L. (1995) New species and new records of anurans from Borneo. The Raffles Bulletin of Zoology, 43 (1), 115 - 131.","Malkmus, R. & Matsui, M. (1997) Megophrys kobayashii ein neuer pelobatider frosch vom Mount Kinabalu. Sauria, 19, 31 - 37. [in Deutsch]","Inger, R. F. & Iskandar, D. T. (2005) A collection of amphibians from West Sumatra, with description of new species of Megophrys (Amphibia: Anura). The Raffles Bulletin of Zoology, 53 (1), 133 - 142."]}

Megophrys lancip sp. nov. (Figs. 3–5) Holotype. MZB Amph 22233 (field number ENS 14509) an adult male from the north-east of Desa Ngarip, Kecamatan Ulubelu, Kabupaten Tanggamus, Province of Lampung, Sumatra (5.31359°S, 104.53151°E, elevation 1010 m. a.s.l.), collected by Eric Nelson Smith, Amir Hamidy and Elijah Wostl, at 2005 h on 10 June 2013 (Fig. 3, 4A–E). Paratypes. UTA A-64763 (field number ENS 7579; formerly MZB Amph 12564) adult male from the Bengkulu to Kepahiang road, Kecamatan Taba penanjung, Kabupaten Bengkulu Tengah, Province of Bengkulu, Sumatra (3.68561°S, 102.53861°E, 620 m. a.s.l.), collected by Eric N. Smith and Michael B. Harvey, at 1740 h on 30 May 1996 (Fig. 5B); MZB Amph 3469, 14602, and 26738 (Fig. 4F–J), three adult females from Bukit Barisan Selatan National Park, at Desa Kubu Perahu, near Kota Liwa, Kecamatan Balik Bukit, Kabupaten Lampung Barat, Province of Lampung, Sumatra, collected by Andiek Fajar on 19 February 1998 (5.07870°S, 104.01189°E, 434 m. a.s.l.), by Inggar Ul-Hasanah on 17 September 2005 (5.07526°S, 104.04081°E, 550 m. a.s.l.), and by Anton Nurcahyo in 2005 (5.07526°S, 104.04081°E, 550 m. a.s.l.), respectively; MZB Amph 3 472 adult female from Bukit Barisan Selatan National Park, at Desa Sukaraja, Kecamatan Semaka, Kabupaten Tanggamus, Province of Lampung, Sumatra (5.51955°S, 104.45303°E, 526 m. a.s.l.), collected by Andiek Fajar on 6 November 1997. Etymology. The species name lancip is the Indonesian word for “pointed”, used as an adjective, and reflects the extremely pointed rostral appendage of the new species. Suggested English common name. Pointed Horned Frog. Suggested Indonesian common name. Katak-tanduk lancip. Diagnosis. The new species is assigned to the genus and subgenus Megophrys based on our molecular analysis (Fig. 2) and the following morphological features: (1) absence of a distinct glandular fold forming a Y, X, or H on the parietoscapular region, possession of (2) a broad and depressed head, (3) pectoral and femoral glands, (4) eyelid appendages, (5) a vertical pupil, (6) a pair of conical spine at the corner of the mouth, and (7) a protruding snout (Delorme et al. 2006). Megophrys lancip sp. nov. is a medium-sized Megophrys (adult males SVLh 37.9–47.7 mm, n = 2; adult females SVLh 38.7–82.5 mm, n = 4) with the head shorter than the body (adult males HL 15.3–19.9 mm: 40.3–41.7% of SVLh, n = 2; adult females HL 16.0– 35.4 mm: 41.4–42.9% of SVLh, n = 4), and broader compared to its length (adult males HL 87.7–95.8% of HW, n = 2; adult females 80.8–85.7% of HW, n = 4). The snout is moderate length (adult males SL 30.5–35.8% of HL, n = 2; adult females SL 29.4–33.6% of HL, n = 4), extremely pointed in dorsal view, protruding in lateral view, and with a sharp and pointed (acuminate) rostral appendage that projects beyond the lower jaw. The rostral appendage is moderate (adult males SNLh 3.0– 3.3 mm, 7.0–8.0% to SVLh, n = 2; adult females SNLh 2.5–5.4 mm, 4.9–6.7% of SVLh, n = 4; adult males SAL; 1.2–1.4 mm, 2.9–3.1% of SVLh, n = 2; adult females SAL 0.6–3.44 mm, 1.5–4.1% of SVLh, n = 4); the upper eyelid has a triangular, short, and narrow-based appendage; the tympanum is distinct, small, oval, andoriented vertically; vomerine teeth are present; and the subarticular tubercle and outer metatarsal tubercles are absent. Description of Holotype (measurements in mm). Adult male; body stocky, robust (SVL 36.8; SVLh 37.9); head broad and depressed (HW 17.4: 46.0% of SVLh), slightly broader than long (HL 15.3: 40.3% of SVLh); snout extremely pointed from dorsal view, protruding in lateral view, tip with an acuminate sharp rostral appendage projection (SAL 1.2: 3.1% of SVLh), projecting beyond lower jaw; eye large and lateral on head, smaller (ED 5.0: 13.1% of SVLh) than snout length (SL 5.5: 14.4% of SVLh), eye diameter about 3.5 times of tympanum diameter (ED 5.0: 359.4% of TDH); canthus rostralis sharp, angular, concave; nostril below canthus on lateral view, distinctly closer to eye (NEL 1.9: 4.9% of SVLh) than to snout (SNL 2.3: 6.0% of SVLh); internarial distance (IND 3.4: 8.9% of SVLh) much shorter than interorbital distance (IOD 5.1: 13.5% of SVLh); upper eyelid with low tubercles and short narrow-base eyelid appendage projection (EHL 1.2: 2.9% of SVLh) much shorter than total width (UEWh 4.8: 12.7% of SVLh); a conical spine present at corner of mouth; tympanum distinct, oval, oriented vertically (TDV 2.2: 5.7% of SVLh; TDH 1.4: 3.6% of SVLh); vomerine teeth present; tongue not notched, without papilla. Forelimbs slender and short (FLL 25.5: 67.3% of SVLh), about half of hindlimbs (FLL 25.5: 49.0% of HLL); fingers moderately slender, unwebbed, without lateral fringes, with rounded and swollen tips; first finger (fin1L 3.1: 8.1% of SVLh) slightly longer than second (fin2L 3.0: 7.8% of SVLh) and fourth (fin4L 2.8: 7.5% of SVLh), third much longer (fin3L 5.0: 13.1% of SVLh); finger length formula IV Skin. Dorsal skin is smooth with low, sharply angular, tubercles from the snout to the vent, especially around the waist; the tubercles are denser laterally on the body and on the posterior surface of the thigh; upper eyelids with low conical tubercles; supratympanic fold distinct, angular, widening anteriorly, narrow posteriorly, extending from posterior of eye, curving down around upper border of tympanum and terminating above axilla; Y, X or H fold absent between parietoscapular area to level of axilla; dorsolateral fold extends from between eye and tympanum to the groin, wider from shoulder to groin; forelimb and hindlimb surface tuberculate and with transverse folds in forearm, tibia, and femur; a weakly developed tubercles ridge on outer of edge of forearm and tarsus forming a weak serration; ventral skin slightly tuberculate; skin on throat wrinkled, chest slightly tuberculate; ventral surface of limbs and abdomen smooth to weakly granular; a pair of white pectoral glands, small, conical and slightly raised; a pair of white femoral glands, moderately large, rounded and slightly raised, at mid-femur. Male with a pair of vocal slits on the posterior corners of the mouth, the single, external, vocal sac is indistinct on the preserved specimen; dark nuptial pads are present on the dorsomedial surface of the first and second fingers, from the base of the finger to the first terminal joint. Colour. In life (Fig. 3A–D), the entire dorsal surface of the head, body, forelimbs and hindlimbs is light brown; a darker brown inverse triangular marking bounded by a thin fold is present from the interorbital region to the posterior area of the head. The dorsal surface of the body, forelimbs and hindlimbs have indistinct, darker brown blotches. The dorsal surface of the lower arms, tibia and femur (hindlimbs) have dark brown transverse folds. The lateral side of the head, including the canthal area, is brown, darker around the nostrils and just posterior and below the canthus, lighter anterior and posterior to the eye, with dark markings above and below the skin bordering the eye; supratympanic fold light brown, with indistinct dark brown blotches. The entire ventral surface is light brown with a heavy dark transverse marking and dark blotches over the throat and on the chest and abdomen. Ventral surface of throat to chest is dark brown, while the abdomen, forelimbs and hindlimbs are light cream, with some small blotches. In preservative (Fig. 4A–E) aspects of the colour pattern remain, but the dorsum darkens. Variation (measurements in mm). Individuals of the type series are morphometrically generally similar (Table 4 and 5). Based on the individual specimens collected, females appear to be slightly larger and stockier than males (male SVL 36.8–46.3 mm [n = 2] vs female SVL 38.2–79.1 mm [n = 4]). When the hindlimb is bent forward along the body, the tibiotarsal joint usually reaches to the posterior corner of the eye in adult males and reaches to the posterior of tympanum in females. Only one female (MZB Amph 3 472) had the tibiotarsal joint reach the posterior of the eye. Individuals of the type series are only slightly variable in colouration. In preservative, the dorsal colouration in both males and females is relatively uniform light brown, but is completely dark brown in the single male paratype (MZB Amph 12564). The colouration of the ventral surface in both sexes is uniformly brown with dark blotches and dark longitudinal bands on the throat and chest. This longitudinal pattern is faded in one male (MZB Amph 12564). Comparisons. Megophrys lancip sp. nov. differs from members of subgenera Xenophrys, Atympanophrys, Panophrys, and M. dringi by the absence of a folded gland forming a Y, X or H directed backwards at the parietoscapular region (Günther, 1864; Delorme et al. 2006) (versus presence) (Figs. 5A, 6C–D, Table 6). Megophrys lancip sp. nov. is distinguished from all known members of Sundaland and Philippine Megophrys (except for M. montana and M. nasuta), by the presence of a well -developed rostral appendage on the tip of the snout (versus absence in M. aceras, M. baluensis, M. dringi, M. edwardinae, M. longipes, M. kobayashii, and M. parallela) (Table 6, Fig. 5B–D). Megophrys lancip sp. nov. differs from Philippine M. stejnegeri by having a triangular-shaped eyelid appendage (versus a tubercle) and from M. ligayae by having a single dorsolateral fold that extends from the parietoscapular region to the groin (Table 6, type I) (versus a dorsolateral folds that extends from the parietoscapular region to the middle of the body [Table 6, type II]). Megophrys lancip sp. nov. can be distinguished from M. nasuta by absence of additional lateral fold that extends from the area behind the supratympanic fold to the groin (versus presence in M. nasuta) (Table 6; Fig. 5D), shorter rostral appendage SAL 1.5–4.1% of SVLh (versus SAL 1.2–9.2% of SVLh), and shorter eyelid appendage EHL 0.8–2.2 mm; 22.4% of UEWh (versus EHL 3.5–11.9 mm; 50.6% of UEWh) (Table 5; Fig. 5B, D) Megophrys lancip sp. nov. differs from M. montana by having a smaller body size, SVL 36.8–46.3 mm in adult males and 38.2–79.1 mm in adult females (versus 38.1–53.9 mm in adult males and 45.7–99.5 mm in adult females) (Table 4), longer snout appendage, SAL 1.5–4.1% of SVLh, (versus SAL 0.0–3.5% of SVLh), shorter eyelid appendage EHL 0.8–2.2 mm; 22.4% of UEWh (versus EHL 0.9–3.9 mm; 30.0% of UEWh), shorter hand length, 8.6–10.4 mm in adult males and 9.8–19.0 mm in adult females, (versus 10.4–14.4 mm in adult males and 11.0– 23.6 mm in adult females), shorter hindlimb, HLL 52.0– 58.8 mm in adult males and 43.4–98.3 mm in adult females, (versus 56.2–76.6 mm in adult males and 56.0– 133.4 mm in adult females), tibio-tarsal articulation reached behind the eye in adult males and mostly reached behind the tympanum in adult females, (versus reached to the front of eye in adult males and reached the middle of eye in adult females), and less developed toe webbing I (1), II (1–1), III (2–2), IV (3–3), V (1 1/2) [versus I (1), II (1–1/2), III (1 3/4–1 1/2), IV (2 1/2–2 1/2), V(1)] (Table 6). TABLE ³. Variation in size anđ bođy proportions of male anđ female Megophrys from the Sunđalanđ anđ Philippines, mođifieđ from Taylor (1920), Inger (1954), Inger et al. (1995), Malkmus & Matsui (1997), anđ Inger & Iskanđar (2005). * for M. parallela, the measurement đoes not incluđe the rostral appenđage. ……continued on the next page ……continued on the next page ……continued on the next page Megophrys lancip sp nov. Megophrys montana Species RSAL RHLh RHW RAG REHL RLAL RHLL RFML RTL REHL/UEWh RTDH/ED RFL/TL RHW/HLh Species male (n = 28) female (n = 21) juvenile (n = 12) male (n =6) female (n = 2) RSAL 4.2 (1.5–8.7) 4.2 (1.8–9.2) 2.4 (1.2–5.4) – – – – RHLh 44.0 (39.5–49.7) 46.0 (40.6–52.8) 42.0 (36.9–46.7) 42.6 (39.1–44.3) 41.8 (41.6–42.0) RHW 46.4 (44.0–49.5) 48.4 (44.8–54.8) 45.2 (37.0–49.4) 46.6 (44.9–51.9) 48.5 (48.5–48.6) RAG 42.0 (33.5–47.0) 46.5 (39.6–57.2) 48.4 (42.7–51.6) 44.5 (40.1–47.3) 46.3 (40.1–52.6) REHL 8.3 (6.3–11.4) 8.9 (4.4–13.0) 9.6 (6.0–12.7) 2.8 (1.8–3.4) 3.7 (3.2–4.2) RLAL 46.6 (20.8–49.8) 32.4 (19.9–48.2) 23.3 (20.7–25.2) 49.3 (46.3–50.5) 47.8 (47.2–48.5) RHLL 121.0 (108.8–147.9) 116.8 (106.6–126.2) 119.0 (111.6–129.0) 148.4 (139.9–161.5) 147.2 (143.0–151.4) RFML 40.5 (36.0–48.1) 38.9 (33.2–41.9) 40.4 (37.5–44.8) 48.1 (44.3–51.0) 47.5 (45.2–49.7) RTL 36.0 (33.3–44.1) 35.1 (31.8–37.9) 37.1 (34.4–40.4) 43.9 (41.3–48.4) 44.4 (42.4–46.4) REHL /UEWh 50 (38.5–61.4) 53.6 (32.7–61.3) 48.4 (32.1–62.7) 21.9 (14.7–29.0) 30.6 (24.9–36.3) RTDH /ED 32.7 (22.2–43.9) 35.7 (16.3–52.8) 28.2 (10.9–34.5) 32.4 (22.1–35.8) 31.5 (29.6–33.3) RFL/TL 111.8 (103.0–118.2) 111.1 (98.8–121.3) 108.9 (101–113.3) 107.1 (105.4–112.5) 106.9 (106.6–107.2) RHW /HLh 106.4 (92.9–116.1) 103.9 (92.4–119.5) 106.8 (91.6–111.2) 111.7 (106.9–118.1) 116.1 (115.4–116.9) Megophrys parallela Relative values (R) of each character compared to SVL (* used HL, and HW compared to HL). * Fold forming Y, X or H on the parietoscapular region to the level of axilla. ** Dorsolateral fold shape: Type I, dorsolateral folds are elongated and extend from the parietoscapular region to the groin. Type II, dorsolateral folds extend from the parietoscapular region to the middle of the body Type III, multiple dorsolateral folds - at least three or four - and they are discontinuous, formed by a series of elongated tubercles. Type IV, dorsolateral folds are elongated and curve from the axillary region towards (and reaching) the posteriodorsal margin of tympanum. Distribution and Natural History. The new species is known from the provinces of Lampung and Bengkulu in southwestern Sumatra (Fig. 1). Larval, acoustic and other ecological data are unknown. The holotype was collected from a coffee plantation near the edge of secondary forest at 1010 m a.s.l. The male paratype from Bengkulu was collected from inside a rotten log on the forest floor at 620 m a.s.l. The female paratypes, collected from Bukit Barisan Selatan National Park, were collected from primary forest between 434 and 526 m a.s.l. This new species of Megophrys can be found sympatrically with M. nasuta in Kubu Prahu, Bukit Barisan Selatan National Park. Habitat loss and exploitation for the pet trade are likely be the main threats for the new species.

KEY TO THE HONDURAN SPECIES OF THE TANTILLA TAENIATA SPECIES GROUP AND THE GUATEMALAN SPECIES T. TAENIATA (AS RESTRICTED TO SOUTHEASTERN GUATEMALA BY CAMPBELL 1998 AND HEREIN) 1. A. Pale vertebral stripe confined to vertebral row throughout its length; maximum known total length more than 500 mm..................................... T. impensa B. Pale vertebral stripe variable; maximum known total length less than 450 mm............................................. 2 2. A. Pale vertebral stripe reduced to series of dashes confined to middle of vertebral row.......... T. olŋmpia B. Pale vertebral stripe complete, restricted to vertebral row, or extending onto edges of paravertebral rows................................................................................... 3 3. A. Pale vertebral stripe confined to vertebral scale row on at least anterior half of body.......... T. stenigrammi B. Pale vertebral stripe extending onto edges of paravertebral scale rows throughout its length................ 4 4. A. Ventral surfaces some shade of red............................ 5 B. Ventral surfaces yellow or white................................ 6 5. A. Ventrals 141–152 in males, 150 in known female.............................................................................. T. taeniata B. Ventrals 153–163 in males, 154–161 in females................................................................................ T. psittaca 6. A. Pale nuchal collar divided; ventrals 157 in one male, 155–161 in two females; máximum known total length 273 mm.................................................... T. tritaeniata B. Pale nuchal collar complete; ventrals variable; máximum known total length more than 300 mm........... 7 7. A. Ventrals 169 in one male, 161–178 in two females............................................................................. T. excelsa B. Ventrals 142–158 in both sexes combined.... T. gottei, Published as part of McCranie, James R. & Smith, Eric N., 2017, A Review of the Tantilla taeniata Species Group (Reptilia: Squamata: Colubridae: Colubrinae) in Honduras, with the Description of Three New Species, pp. 338-348 in Herpetologica 73 (4) on page 347, DOI: 10.1655/Herpetologica-D-16-00080.1, http://zenodo.org/record/7712317, {"references":["Campbell, J. A. 1998. Comments on the identities of certain Tantilla (Squamata: Colubridae) from Guatemala, with the descriptions of two new species. Scientific Papers, Natural History Museum, University of Kansas 7: 1 - 14."]}

Tantilla excelsa sp. nov. (Figs. 6, 7) Tantilla taeniata: Wilson and Meyer 1971:32, in part; Wilson 1982:56, in part; Wilson and Meyer 1982:109, in part; Wilson and Meyer 1985:103, in part; Wilson and McCranie 1999:328, in part; McCranie 2011a:227, in part; McCranie 2011b:44, in part; McCranie et al. 2012:623; Wilson and Mata-Silva 2015:457, in part. Holotype. — USNM 579682, an adult male from Lancetilla (15°44 ′ N, 87°27 ′ W), 30 m elevation, department of Atlántida, Honduras, collected 26 February 2011 by Leonel Marineros. Paratypes (n ¼ 3). — UF 157566, an adult female from Compañia Agricola Paradise near Peña Blanca (14°58 ′ N, 88°01 ′ W), 700 m elevation, Cortés, Honduras; UMMZ 58417, an adult female that is dehydrated and has an incomplete tail, from El Progreso (15°24 ′ N, 87°48 ′ W), Yoro, Honduras, 30 m elevation; MCZ 22045, head and anterior portion of body of an adult, from Progreso District, Yoro, Honduras, about 30 m elevation. Diagnosis. — Tantilla excelsa (in preservative) is defined by the following combination of characters: (1) pale middorsal stripe tan, extending length of body and at least half of tail, stripe on vertebral row and adjacent halves of paravertebral rows throughout its length; (2) tan to white lateral stripe occupying adjacent halves of scale rows 3 and 4; (3) ventrolateral area similar in pale color to that of lateral stripe; (4) lower two-thirds anteriorly and about lower third posteriorly of scale row 1 white similar to that of ventrals; (5) lateral edges of ventrals white, no, or very few, dark spots present; (6) ventral and subcaudal surfaces yellow (shortly after death) and white; (7) two females with 161–178 ventrals, single male with 169 ventrals, subcaudals 61 in one female, 70 in male; (8) tail length 23% of total length in one female, 24% in male. Tantilla excelsa can be distinguished from T. taeniata by having 169 ventrals in single male, 161–178 in two females, and by having yellow ventral surfaces in life (vs. 141–152 ventrals in males and 150 in single known female, and ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and tail). Tantilla excelsa differs from T. impensa and T. stenigrammi by having the middorsal pale stripe involving all of the vertebral scale row and adjacent thirds of paravertebral scale rows throughout its length (vs. middorsal stripe confined to vertebral row in those two species [on at least anterior half of body in T. stenigrammi]) and having female ventrals numbering up to 178 (vs. to 172 in T. impensa and 159 in female T. stenigrammi). Tantilla excelsa differs from T. olŋmpia in having complete middorsal and lateral pale stripes, with the middorsal stripe extending onto lateral edges of paravertebral rows, and having 169 ventrals and 70 subcaudal scales in single male (vs. those stripes reduced to dashes and spots, respectively; pale middorsal dashes restricted to vertebral row; and 148 ventrals and 49 subcaudals in a male specimen). Tantilla excelsa differs from T. psittaca by having yellow ventral and subcaudal surfaces in life and white to pale yellow in preservative and in having 169 ventrals in single male and 161–178 in two females (vs. those surfaces pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative, 153–163 ventrals in males and 154–161 in female specimens). Tantilla excelsa differs from T. tritaeniata in having 169 ventrals in single male and 161–178 ventrals in two females, and having a complete pale nuchal collar (vs. 157 ventrals in single male and 155–161 in females, and pale nuchal collar divided). Description of holotype. —An adult male; TOL 400 mm; SVL 305 mm; TAL 95 mm (23.8% of TOL); HL 9.8 mm; HW 5.5 mm (level of angle of mouth); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.3 mm; snout length 3.0 mm, about 2.3 times longer than eye; pupil circular; rostral about 1.1 times wider than high (2.2 X 2.0 mm); internasal length about 0.8 times width (1.1 X 1.4 mm); prefrontal larger than internasal, wider than long (2.2 X 1.5 mm); median prefrontal suture 1.4 mm, about half as long as frontal; frontal with slight anterior extension, Vshaped posteriorly, about 1.0 times longer than wide (2.9 X 2.8 mm), about 1.0 times longer than distance from its anterior edge to tip of snout (2.9 X 2.8 mm); parietal about 1.5 times longer than wide (4.1 X 2.8 mm), median parietal suture length 3.0 mm, about 1.0 times longer than length of frontal (2.9 mm); parietals contacting five nuchal scales; supraocular about 1.6 times longer than wide (2.3 X 1.4 mm), bordering orbit, contacting parietal, upper postocular, preocular, frontal, and prefrontal. Nasal divided, anterior nasal contacting rostral, internasal, and supralabial 1, posterior nasal contacting internasal, prefrontal, preocular, and supralabials 1 and 2, nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, about 1.4 times higher than long (0.7 m X 0.5 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about 1.5 times larger than lower (upper height 0.6 mm, lower height 0.4 mm); temporals 1 þ 1, anterior temporal 1.5 times longer than high (1.9 X 1.3 mm), posterior temporal 1.8 times longer than high (1.6 X 0.9 mm); supralabials 7, 3 and 4 bordering orbit, 4 and 5 contacting lower postocular, 5 also contacting anterior temporal, 6 contacting anterior temporal, 7 contacting anterior and posterior temporals; infralabials 6– 7, first 2 contacting anterior pair of chinshields on side with 6 infralabials (left side), first 3 contacting anterior pair of chinshields on side with 7 infralabials (right side); infralabial 1 on each side separated medially by contact with anterior chinshield; mental about 1.6 times wider than long (1.4 X 0.9 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields about 2.4 times longer than wide (2.6 X 1.1 mm), not extending laterally to border of lip; posterior pair of chinshields about 1.7 times longer than wide (1.7 X 1.0 mm), posterior tips separated from each other by two rows of gular scales; four preventral scales between gular 1 and ventral 1; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in six rows at level of subcaudal 10; ventrals 169; cloacal scute divided; subcaudals 70, paired; ventrals plus subcaudals 239. Color of holotype in life. —Other than noting the ventral and subcaudal surfaces were yellow (L. Marineros, personal communication), the collector of the holotype did not record color notes of the recently killed specimen. Color of holotype in preservative (Figs. 6, 7). —Dorsal surfaces brown with white nuchal collar, tan middorsal stripe, and white lateral stripe; tan middorsal stripe includes vertebral row and adjacent tips to a third of paravertebral scale rows; tan middorsal stripe narrowing anteriorly on first scale involved in stripe, extending onto tail, fading out at about midlength of tail; tan middorsal stripe beginning on scale 4 posterior to parietals; broader white lateral stripe present, covering adjacent halves of scale rows 3 and 4, bordered above by continuous, thin dark brown line; white lateral stripe beginning on scale 5 posterior to supralabial 7, extending well onto tail to nearly reaching tip; lower twothirds of scale row 1 white anteriorly, grading to white only reaching lower tips of scale row 1 posteriorly on body; about upper third of scale row 1, all of scale row 2, and lower third of scale row 3 same shade of brown as paravertebral rows on about anterior half of body, before reducing width posteriorly; pale nuchal collar complete, not involving posterior tips of parietals, but involving first two scale rows dorsally, pale nuchal collar also involving posterior half of ultimate supralabial plus two scale rows laterally; dorsal surface of head brown, other than dark brown line crossing posterior tips of parietals to connect with lateral dark head bar; white, with dirty white flecking lateral spot involving postnasal, supralabial 1, and anterior half of supralabial 2; larger white spot posterior to eye involving anterior half of anterior temporal, all of supralabial 5, and tips of supralabials 4 and 6; dark brown lateral bar involving posterior half of supralabial 7, adjacent two-thirds of supralabial 6, and all of posterior temporal to connect with dark brown parietal cross-line; lateral dark bar narrowing only slightly dorsally; anterior tip of mental scale and lateral tips of infralabials 1 and 2 with brown flecking; dark brown spot present on infralabial 4 and adjacent edge of infralabial 5, remainder of ventral surface of head white; ventral surfaces of body and tail white for full length, except for variation discussed above. Variation in paratypes. —Two adult female paratypes show the following variation (followed by mean in parentheses): TOL 326 mm in one (UF 157566; UMMZ 58417 has incomplete tail); SVL 251–276 (263.5). mm; TAL 23.0% of TOL in one; ventrals 161–178 (169.5); subcaudals 61 in one; ventrals plus subcaudals 222 in one. All paratypes (including MCZ R22045, a head and anterior portion of body only of an adult) have 7 supralabials on each side, with supralabials 3 and 4 bordering the eye. Available color pattern notes indicate those paratypes were similar overall to the color pattern of the holotype. Distribution and habitat. —Very little is known about the habitats of Tantilla excelsa. The holotype was killed by a worker at the Lancetilla Botanical Gardens during February. The UF specimen was taken after dark as it was crawling on the ground on a coffee farm during May. The UMMZ specimen was collected on 27 March 1923 and possibly came from an insect trap (G. Schneider, personal communication). No data are available for the MCZ paratype other than it was collected in 1925. The Lancetilla Botanical Gardens formerly contained Lowland Moist Forest and lies at 30 m elevation. The coffee farm is in an area formerly containing Premontane Wet Forest and lies at 700 m elevation. The remaining two localities are in Lowland Dry Forest, although more mesic corridors of gallery forest do occur in those areas. Thus, T. excelsa is known to occur at low and moderate elevations (30–700 m) on both sides of the Río Ulúa drainage, and from the vicinity of Lago de Yojoa, Cortés, to the north-northeast in the vicinity of Lancetilla, Atlántida (Fig. 5). Etymology. —The specific name excelsa is an adjective from the Latin excelsus, which in this case means high. The name refers to the relatively high number of ventral scales in this nominal form.

Tantilla stenigrammi sp. nov. (Figs. 3, 4) Tantilla taeniata: McCranie 2011a:227, in part; McCranie 2011b:44, in part. Holotype. — University of Texas at Arlington (UTA) R-52591 (Field No. ENS 10897), a subadult female from Cuaca (15°23 ′ 01.75 ′′ N, 86°12 ′ 41.22 ′′ W;. datum ¼ WGS 84 in all cases), 895 m elevation, department of Olancho, Honduras, collected 3 February 2005 by Eric N Smith, Jorge A. FerrariCastro, Jorge Murillo, C. Chavez, Arturo Sosa, and John H. Malone. Paratype (n ¼ 1). — UNAH 3032, an adult male with an incomplete head and tail, from about 20 km E of Gualaco, Olancho, Honduras, at beginning of a foot trail (15°02 ′ N, 85°54 ′ W) leading to Cerro La Picucha, 1180 m elevation. Diagnosis. — Tantilla stenigrammi is defined, following Campbell (1998), by the following combination of characters: (1) middorsal stripe Warm Buff to Pale Horn Color (Color 118, 92), extending length of body and most of tail, confined to vertebral row on at least anterior half of body, extending onto adjacent edges of paravertebral scale rows posteriorly on body; (2) Buff-Yellow (Color 53) lateral stripe occupying adjacent thirds of scale rows 3 and 4; (3) ventrolateral area similar in pale color to that of lateral stripe; (4) lower half to two-thirds of scale row 1 colored similarly to that of ventrals; (5) lateral edges of ventrals immaculate white (in preservative), no dark spots present; (6) ventral and subcaudal surfaces Pearl Gray (in life; Color 81) to white (in preservative); (7) female with 159 ventrals, male with 164 ventrals; (8) tail incomplete in both. Tantilla stenigrammi can be distinguished from T. taeniata by having the pale middorsal stripe confined to the vertebral row on at least anterior half of body and in having pale gray ventral surfaces in life (vs. pale middorsal stripe also extending onto paravertebral rows throughout its length and ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and under tail in T. taeniata). It further differs from T. taeniata in having more ventral scales (164 vs. 141–152 in males, and 159 vs. 150 in females). Tantilla stenigrammi differs from T. psittaca and T. tritaeniata by having the middorsal pale stripe confined to the vertebral row on at least anterior half of body (vs. middorsal stripe involving all of vertebral scale. and adjacent third to half of paravertebral scale rows throughout its length in those two species); further from T psittaca by having pale gray ventral and subcaudal surfaces in life and white in preservative (vs. those surfaces being pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative), and further from T. tritaeniata in having a complete nuchal collar and the pale lateral stripe covering adjacent thirds of scale rows 3 and 4 (vs. nuchal collar divided and pale lateral stripe including adjacent halves to two-thirds of scale rows 3 and 4). Tantilla stenigrammi differs from T. impensa in having 159 ventrals in the female, the pale middorsal stripe extending onto adjacent edges of paravertebral scale rows only posteriorly on the body, the pale brown lateral stripe covering adjacent thirds of scale rows 3 and 4, and by having a pale brown crossbar on the internasals that is bordered on the posterior edge by a dark brown cross-line (vs. 164–172 ventrals in females, the pale middorsal stripe confined to vertebral row throughout its length, pale lateral stripe including adjacent halves to twothirds of scale rows 3 and 4, and those head surfaces same ground color as remainder of head). Tantilla stenigrammi differs from T. olŋmpia in having complete middorsal and lateral pale stripes, in having a pale internasal crossbar, and having distinct pale large spots on the lateral surface of the head (vs. those body pale stripes reduced to dashes and spots, respectively, no internasal crossbar, and those lateral pale areas on head absent to indistinct). Description of holotype. —A subadult female; TOL 173 mm plus tail tip; SVL 137 mm; TAL 36 mm plus tail tip (20.8% plus tail tip of TOL); HL 4.6 mm; HW 3.4 mm (level of angle of mouth); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.1 mm; snout length 2.2 mm, about 2.0 times longer than eye; pupil circular; rostral about 1.2 times wider than high (1.2 X 1.0 mm); internasal length about three-quarters of width (0.8 X 1.1 mm); prefrontal much larger than internasal, wider than long (1.3 X 0.9 mm); median prefrontal suture 0.9 mm, nearly half as long as frontal; frontal with slight anterior extension, V-shaped posteriorly, about 1.3 times longer than wide (2.1 X 1.6 mm), 1.3 times longer than distance from its anterior edge to tip of snout (2.1 X 1.6 mm); parietal about 1.6 times longer than wide (3.0 X 1.9 mm), median suture length 1.6 mm, about 0.8 times frontal length; parietals contacting seven nuchal scales; supraocular about 1.8 times longer than wide (1.6 X 0.9 mm), bordering orbit, contacting parietal, upper postocular, preocular, and prefrontal. Nasal divided, anterior nasal contacting rostral, internasal, and first supralabial, posterior nasal contacting internasal, prefrontal, preocular, and first and second supralabials, nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, as high as long (both dimensions 0.5 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about same size as lower (height of both 0.5 mm); temporals 1 þ 1, anterior temporal 1.1 times longer than high (1.0 X 0.9 mm), posterior temporal 1.6 times longer than high (1.3 X 0.8 mm); supralabials 7, 3rd and 4th bordering orbit, 4th and 5th contacting lower postocular, 5th and 6th contacting anterior temporal, and 7th contacting anterior and posterior temporals; infralabials 6, first 3 contacting anterior pair of chinshields, infralabial 1 on each side separated medially by contact with anterior chinshield; mental 1.7 times wider than long (1.2 X 0.7 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields 2.1 times longer than wide (1.9 X 0.9 mm), not extending laterally to border of lip; posterior pair of chinshields 1.9 times longer than wide (1.3 X 0.7 mm), posterior tips separated from each other by one gular scale; three preventral scales present between gular and ventral 1; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in eight rows at level of subcaudal 10; ventrals 159; cloacal scute divided; subcaudals 64 plus tail tip (an estimated 5–10 subcaudal scales missing), paired; ventrals plus subcaudals 223 plus tail tip. Ventrals Subcaudals TAL/TOL n SpeciesMales/FemalesMalesFemalesMalesFemalesMalesFemalesMaximum TOL T. excelsa 3/1169161–178706124%23%400 mm T. gottei 6/1142–15814762–677024–26%26%367 mm T. impensa 9/12161–171164–17268–7264–7221–24%21–25% ~ 725 mm T. olŋmpia 1/0148—49—21%—338 mm T. psittaca 4/3153–163154–16163–73—24–25%—413 mm T. stenigrammi 1/1164159————173 mmþ T. taeniata 9/1141–15215060–705923–27%23%415 mm T. tritaeniata 1/2157155–161—59–65—23–24%273 mm Color of holotype in life (Fig. 3). —Coloration of freshly euthanized holotype (UTADC 8752–53): dorsum of head Olive-Brown (28), transforming to Sepia (219) along scale margins; anterior portion of rostral, supralabial 1, and nasals Sepia; posterior portion of rostral, internasal, and interparietal areas Grayish Horn Color (91); two Pale Horn Color (92) lateral spots on head, first occupying postnasal, lower edge of prefrontal, anterior tip of preocular, and most of supralabials 1 and 2, second lateral spot located posterior to eye and occupying most of lower postocular, lower borders of upper postocular and parietal, part of supralabials 4 and 6 and all of supralabial 5, and anterior half of anterior temporal; nuchal collar Warm Buff (118) dorsally to Pale Horn Color laterally, with Sepia anterior and posterior borders, collar about two dorsal scales long and occupying posterior tips of parietal and secondary temporal, and posterior third of supralabial 7; dorsum of body Hair Brown (119A), turning Sepia alongside all pale body stripes and toward nuchal collar; middorsal stripe Warm Buff, occupying only vertebral scale row anteriorly on body, extending onto adjacent edges of paravertebral scale rows posteriorly on body, beginning one and a half scales posterior to nuchal collar, stripe extending onto tail to reach tip, occupying adjacent halves of two most-middorsal scale rows; lateral body stripe on each side Buff-Yellow (53), occupying adjacent thirds of scale rows 3 and 4, beginning three scales posterior to nuchal collar, pale stripe on tail occupying second dorsal scale row and ending about 10 scales anterior to tip; underside of head mostly Pearl Gray (81), but Smoke Gray (44) toward mouth, and mental and buccal borders of infralabials 1–4 Sepia; venter of body, including anal plate, Pearl Gray, Smoke Gray toward sides, underside of tail Pearl Gray; ventral coloration extends to middle of dorsal scale row 1 on body, occupying all of dorsal scale row 1 on tail. Color of holotype in preservative (Fig. 4). —Dorsal surfaces brown with white nuchal collar and white middorsal and lateral stripes; pale middorsal stripe confined to vertebral row on at least anterior half of body, and extending onto adjacent edges of paravertebral rows posteriorly on body, with lateral tips of each vertebral scale dark brown; pale middorsal stripe not narrowing anteriorly on first scale involved in stripe; pale middorsal stripe extending onto tail, fading out at about a third of its length until becoming poorly visible at about midlength of tail; pale middorsal stripe beginning on 4th scale posterior to parietals; pale lateral stripe present on adjacent thirds of scale rows 3 and 4, bordered above by continuous, thin dark brown line; pale lateral stripe beginning on 5th scale posterior to ultimate (7th) supralabial, extending well onto tail; lower two-thirds of scale row 1 white; upper third of scale row 1, all of scale row 2, and lower third of scale row 3 same shade of brown as paravertebral rows; pale nuchal collar complete, involving posterior tips of parietals plus one and one-half scale rows dorsally, pale nuchal collar also involving posterior third of ultimate supralabial plus one scale row laterally; dorsal surface of head generally brown; off-white crossbar involving upper tip of rostral and both internasals, crossbar bordered posteriorly by dark brown cross-line involving posterior tips of internasal scales; lateral tips of each parietal scale white, contacting off-white line following parietal suture; anterior lateral white spot involving lower tips of prefrontals, postnasal, posterior two-thirds of supralabial 1, and all but posterior edge of supralabial 2; larger lateral white spot also present posterior to eye, involving all of supralabial 5 and anterior edge of supralabial 6, extending dorsally to cover about anterior half of anterior temporal; dark brown lateral bar covering posterior half of anterior temporal, all of posterior temporal, and posterior three-quarters and anterior two-thirds of supralabials 6 and 7, respectively; dark brown lateral bar extending dorsally to lateral third of parietal, not narrowing dorsally; about anterior half of mental and lateral tips of infralabials 1 and 2 dark brown, small portions of infralabials 3 and 4 with brown spots, remainder of ventral surface of head white; ventral surface of body white for full length, white pigment also extending onto lower tips of scale row 1 on body; subcaudal surface white, except upper tips of subcaudal scales dark brown. Variation in paratype. —The paratype is an adult male that lacks part of its head and most of its tail, thus no measurements were taken, nor was the color pattern recorded by JRM when examined. Data and photographs in preservative provided by J.E. Mérida (26 December 2016), however, indicate that it has 164 ventrals and a narrow vertebral stripe anteriorly on the body and extending onto adjacent edges of paravertebral scale rows posteriorly on body; thus, similar to that of the holotype. Distribution and habitat. —The holotype of Tantilla stenigrammi was found at 1630 h beside the school in Cuaca, Olancho (Fig. 5). The specimen was under a root mat on the buttress of a fallen tree about 1 m above ground, and that site located between a soccer field and secondary vegetation. The historic habitat of Cuaca was pine–oak forest (Premontane Moist Forest of Holdridge [1967]) along the flanks of Montaña de Botaderos in a transition zone between pine– oak forest and broadleaf rainforest (Premontane Wet Forest). The highest elevations of the Botaderos Mountains contain remnants of closed-canopy broadleaf rainforest (Lower Montane Wet Forest), usually referred to as ‘‘cloud forest.’’ The paratype was found dead during the day near the start of a foot trail in pine–oak forest (Premontane Moist Forest) along the lower flanks of the Sierra de Agalta. That trail also passes through Premontane Wet Forest, before ultimately entering Lower Montane Wet Forest as it gains altitude. Thus, both specimens of Tantilla stenigrammi were found in similar, formerly forested areas, and at similar elevations (895 and 1180 m) in north-central and central Olancho in east-central Honduras. Both localities are in low mountains on both sides of the upper drainage of the Río Sico Tinto (also called Río Negro) that forms the Agalta Valley (Fig. 5). Remarks. —A photograph of the holotype (UTA R-52591) of Tantilla stenigrammi was previously published in McCranie et al. (2006; as T. taeniata). Etymology. —The specific epithet, stenigrammi, is a Greek substantive used as an adjective and meaning narrow-lined, derived from the Greek adjective stenos (narrow) and the Greek noun grammi (line), in allusion to the narrow middorsal stripe that is characteristic of the species., Published as part of McCranie, James R. & Smith, Eric N., 2017, A Review of the Tantilla taeniata Species Group (Reptilia: Squamata: Colubridae: Colubrinae) in Honduras, with the Description of Three New Species, pp. 338-348 in Herpetologica 73 (4) on pages 339-342, DOI: 10.1655/Herpetologica-D-16-00080.1, http://zenodo.org/record/7712317, {"references":["McCranie, J. R. 2011 a. The Snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, USA.","McCranie, J. R. 2011 b. A new species of Tantilla of the taeniata species group (Reptilia, Squamata, Colubridae, Colubrinae) from northeastern Honduras. Zootaxa 3037: 37 - 44.","Campbell, J. A. 1998. Comments on the identities of certain Tantilla (Squamata: Colubridae) from Guatemala, with the descriptions of two new species. Scientific Papers, Natural History Museum, University of Kansas 7: 1 - 14.","Holdridge, L. R. 1967. Life Zone Ecology, revised edition. Tropical Science Center, Costa Rica.","McCranie, J. R., J. H. Townsend, and L. D. Wilson. 2006. The Amphibians and Reptiles of the Honduran Mosquitia. Krieger Publishing Company, USA."]}

Tantilla gottei sp. nov. (Figs. 8, 9) Tantilla taeniata: Wilson and Meyer 1971:32, in part; Wilson 1982:56, in part; Wilson and Meyer 1982:109, in part; Wilson and Meyer 1985:103, in part; Wilson and McCranie 1999:328, in part; McCranie 2011a:227, in part; McCranie 2011b:44, in part; Wilson and Mata-Silva 2015:457, in part. Holotype. — ROM 19996, an adult female from El Picacho (14°07 ′ N, 87°11 ′ W), a zoological park located near Tegucigalpa, 1280 m elevation, department of Francisco Morazán, Honduras, collected 12 December 1986 by Jorge Porras. Paratypes (n ¼ 4). — UNAH 5394, an adult male from Galeras (13°55 ′ N, 86°59 ′ W) about 5 km north of Güinope, 860 m elevation, El Paraíso, Honduras; MCZ R49886, a juvenile male from El Zamorano (14°00 ′ N, 87°01 ′ W), 800 m elevation, Francisco Morazán, Honduras; UNAH 3903, 4833, a subadult male and an adult male, respectively, from Tegucigalpa (centered at about 14°06 ′ N, 87°12 ′ W), 930– 1000 m elevation, Francisco Morazán, Honduras. Referred specimens (n ¼ 2). — UNAH 6141, a dehydrated specimen, from Tegucigalpa (centered at about 14°06 ′ N, 87°12 ′ W), 930–1000 m elevation, Francisco Morazán, Honduras; UNAH 1765 from Alauca (13°51 ′ N, 86°41 ′ W), 500 m elevation, El Paraíso, Honduras. Diagnosis. — Tantilla gottei is defined by the following combination of characters: (1) middorsal stripe pale cream (in preservative), extending length of body and most of tail, on vertebral row and adjacent third of paravertebral rows; (2) pale cream (in preservative) lateral stripe occupying adjacent third of scale row 3 and lower three-quarters of scale row 4; (3) ventrolateral area white as is that of lateral stripe (in preservative); (4) lower two-thirds of scale row 1 colored similarly to that of ventrals; (5) lateral edges of ventrals white, except some dark spots present (in preservative); (6) ventral and subcaudal surfaces yellow (in life) and white (in preservative); (7) female with 147 ventrals, male ventrals 142–158, subcaudals 70 in female, 62–67 in males; (8) female tail length 26% of total length, 24–26% in males. Tantilla gottei most closely resembles T. taeniata in having low numbers of ventral scales, but differs in having yellow ventral surfaces in life, and by having a pale brown internasal crossbar that is confluent with the pale lateral spot anterior to the eye, (vs. ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and tail in life, and in lacking a internasal crossbar). Tantilla gottei differs from T. impensa, T. olŋmpia, and T. stenigrammi by having the middorsal pale stripe involving all of vertebral scale row and adjacent third to half of paravertebral scale rows (vs. middorsal stripe confined to vertebral row in those three species, with that of T. olŋmpia also reduced to a series of dashes; but that stripe also extends onto adjacent edges of paravertebral scale rows posteriorly on body in T. stenigrammi). Tantilla gottei differs further from T. impensa and T. stenigrammi in having 142–152 ventrals in both sexes combined (vs. ventrals 161–172 in T. impensa, and 159 in one T. stenigrammi), and further from T. impensa in having 204–217 ventral plus subcaudal scales in both sexes combined (vs. 233–240). Tantilla gottei differs further from T. olŋmpia in having 62–70 subcaudals in both sexes combined and having a complete pale lateral stripe (vs. 49 subcaudals and pale lateral stripe reduced to dots in T. olŋmpia). Tantilla gottei differs from T. excelsa in having 142–152 ventrals and 204–217 ventral plus subcaudal scales in both sexes combined and having a pale brown internasal crossbar that is confluent with the pale lateral spot anterior to the eye (vs. 161–178 ventrals and 222–239 ventral plus subcaudal scales, and internasal crossbar absent). Tantilla gottei differs from T. psittaca by having yellow ventral and subcaudal surfaces in life and cream in preservative and in having 142–152 ventrals and 204–217 ventrals plus subcaudals in both sexes combined (vs. those surfaces pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative, and 153–163 ventrals and 216–234 ventrals plus subcaudals). Tantilla gottei differs from T. tritaeniata in having 142–152 ventrals in both sexes combined and having a complete pale nuchal collar (vs. 155–161 ventrals and pale nuchal collar divided). Description of holotype. —A subadult female; TOL 367 mm; SVL 270 mm; TAL 97 mm (26.4% of TOL); HL 10.8 mm; HW 5.6 mm (between level of angle of mouth and eye); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.4 mm; snout length 3.3 mm, about 2.4 times longer than eye; pupil circular; rostral about 1.4 times wider than high (2.2 X 1.6 mm); internasal length about 0.7 times of width (1.1 X 1.6 mm); prefrontal about 1.7 times larger than internasal (1.9 X 1.1 mm), as wide as long (both dimensions 1.9 mm); median prefrontal suture 1.6 mm, about 0.5 times as long as frontal (1.6 X 3.5 mm); frontal with slight anterior extension, V-shaped posteriorly, about 1.3 times longer than wide (3.5 X 2.7 mm), about 1.1 times longer than distance from its anterior edge to tip of snout (3.1 mm); parietal about 1.5 times longer than wide (4.4 X 3.0 mm), median suture length 2.8 mm, about 0.8 times length of frontal; parietals contacting six nuchal scales; supraocular about 1.9 times longer than wide (2.6 X 1.4 mm), bordering orbit, contacting parietal, upper postocular, preocular, and prefrontal. Nasal divided; anterior nasal contacting rostral, internasal, and supralabial 1; posterior nasal contacting internasal, prefrontal, preocular, and supralabials 1 and 2; nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, not higher than long (1.3 X 1.4 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about same size as lower (height of both scales 0.9 mm); temporals 1 þ 1, anterior temporal 1.3 times longer than high (1.8 X 1.4 mm), posterior temporal 1.3 times longer than high (1.9 X 1.5 mm); supralabials 7, 3 and 4 bordering orbit, 4 and 5 contacting lower postocular, 5 contacting anterior temporal and lower postocular, 6 contacting anterior temporal, and 7 contacting anterior and posterior temporals; infralabials 6, 1–4 contacting anterior pair of chinshields, first infralabial on each side separated medially by contact with anterior chinshield; mental about 0.9 times as wide as long (1.8 X 1.9 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields about 1.6 times longer than wide (2.2 X 1.4 mm), not extending laterally to border of lip; posterior pair of chinshields about 1.7 times longer than wide (1.7 X 1.0 mm), posterior tips separated from each other by one gular scale; four preventral scales present between gular and first ventral; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in six rows at level of 10th subcaudal; ventrals 147; cloacal scute divided; subcaudals 70, paired; ventrals plus subcaudals 217. Color of holotype in life. —Nothing was recorded on color in life of the holotype, other than that the ventral and subcaudal surfaces were yellow (J. Porras, personal communication). Color of holotype in preservative (Figs. 8, 9). —Dorsal surfaces brown with pale cream nuchal collar and pale cream middorsal and white lateral stripes; pale middorsal stripe on vertebral row and adjacent third of paravertebral rows; pale middorsal stripe slightly narrowing anteriorly on first scale involved in stripe; pale middorsal stripe extending onto tail to nearly its tip; pale middorsal stripe beginning on fifth scale posterior to parietals; pale lateral stripe present on upper two-thirds of scale row 3 and lower three-quarters of scale row 4, bordered above by continuous, thin dark brown line on upper quarter of scale row four and below by incomplete dark brown line on lower third of scale row 3; pale lateral stripe beginning on scale 5 posterior to supralabial 7, extending well onto tail; lower two-thirds of scale row 1 cream; upper third of scale row 1 and all of scale row 2 same shade of brown as dorsal ground color; pale nuchal collar complete, involving posterior tips of parietals plus ~ 1.5 scale rows dorsally, pale nuchal collar also involving posterior third of ultimate supralabial plus one scale row laterally; dorsal surface of head generally brown, except distinct pale brown crossbar involving most of internasals and anterior third of prefrontals, pale crossbar confluent with pale cream lateral spot anterior to eye; lateral pale spot involving postnasal, supralabial 1, and anterior third of supralabial 2; larger pale cream spot also present posterior to eye, involving all of supralabial 5, anterior edge of supralabial 6, and posterior half of supralabial 4, pale spot extending dorsally to cover about anterior half of anterior temporal, and lower postocular; dark brown lateral bar covering posterior twothirds of anterior temporal, all of posterior temporal, and posterior three-quarters and anterior two-thirds of supralabials 6 and 7, respectively; dark brown lateral bar continuous with dorsal surface of head color; dark brown mottling and tiny spots present on mental and first infralabial, respectively; dark brown spot also present on anterior tips of infralabial four; ventral surface of head pale cream; ventral surface of body cream for full length, pale pigment also extending onto lower tips of scale row 1 on body; subcaudal surface cream, except upper tips of subcaudal scales dark brown. Variation in paratypes. —The four male paratypes show the following variation: TOL 165–343 mm (mean ± 1 SD, 265.8 ± 75.1 mm); SVL 125–254 mm (199.5 ± 54.9 mm); TAL 24–26% of TOL; ventrals 142–158 (147.3 ± 4.1); subcaudals 62–67 (64.0 ± 2.2); and ventrals plus subcaudals 204–215 (211.3 ± 5.0). Some of the paratypes were recorded to have slightly narrower pale lateral stripes than the holotype, with those variations involving the adjacent thirds of scale rows 3 and 4. Distribution and habitat. —Almost nothing is known about the habitats of Tantilla gottei, except the low montane habitats where this species was collected are in pine forest in the south-central portion of Honduras (Premontane Moist Forest of Holdridge 1967), with one exception. That exception is a single locality in Lowland Dry Forest (Holdridge 1967). The holotype was collected alive by a worker on the grounds of the El Picacho Zoo during December. One paratype (MCZ R49886) was collected during May. The known elevational range is 500–1280 m. The known geographical distribution of Tantilla gottei is from the upper Río Choluteca drainage, in the vicinity of the capital city of Tegucigalpa, Francisco Morazán, east- and southeast-ward to the south-central portion of the department of El Paraíso (Fig. 5) along the middle portion of the Río Choluteca. Anthropogenic alteration of forest habitats in recent times has further isolated these localities. In addition, most of those surviving pine forests are burned on an annual basis. Those repeated burnings serve to have a permanent drying effect on those burned areas. Therefore, T. gottei should be considered an endangered species. Etymology. —The name gottei is a patronymic noun honoring Steve W. Gotte, a long-time friend of the first author, who also made several field trips with JRM to Honduras. Steve has also been a long-term employee of the US Department of Interior at the National Museum of Natural History in Washington, DC, and most recently in Suitland, Maryland., Published as part of McCranie, James R. & Smith, Eric N., 2017, A Review of the Tantilla taeniata Species Group (Reptilia: Squamata: Colubridae: Colubrinae) in Honduras, with the Description of Three New Species, pp. 338-348 in Herpetologica 73 (4) on pages 344-346, DOI: 10.1655/Herpetologica-D-16-00080.1, http://zenodo.org/record/7712317, {"references":["Wilson, L. D., and J. R. Meyer. 1971. A revision of the taeniata group of the colubrid snake genus Tantilla. Herpetologica 27: 11 - 40.","Wilson, L. D., and J. R. Meyer 1982. The snakes of Honduras. Milwaukee Public Museum Contributions in Biology and Geology 6: 1 - 159.","Wilson, L. D., and J. R. Meyer. 1985. The Snakes of Honduras, 2 nd edition. Milwaukee Public Museum, USA.","Wilson, L. D., and J. R. McCranie. 1999. The systematic status of Honduran populations of the Tantilla taeniata group (Serpentes: Colubridae), with notes on other populations. Amphibia-Reptilia 20: 326 - 329.","McCranie, J. R. 2011 a. The Snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, USA.","McCranie, J. R. 2011 b. A new species of Tantilla of the taeniata species group (Reptilia, Squamata, Colubridae, Colubrinae) from northeastern Honduras. Zootaxa 3037: 37 - 44.","Wilson, L. D., and V. Mata-Silva. 2015. A checklist and key to the snakes of the Tantilla clade (Squamata: Colubridae), with comments on taxonomy, distribution, and conservation. Mesoamerican Herpetology 2: 418 - 498.","Holdridge, L. R. 1967. Life Zone Ecology, revised edition. Tropical Science Center, Costa Rica."]}

Tantilla excelsa sp. nov. (Figs. 6, 7) Tantilla taeniata: Wilson and Meyer 1971:32, in part; Wilson 1982:56, in part; Wilson and Meyer 1982:109, in part; Wilson and Meyer 1985:103, in part; Wilson and McCranie 1999:328, in part; McCranie 2011a:227, in part; McCranie 2011b:44, in part; McCranie et al. 2012:623; Wilson and Mata-Silva 2015:457, in part. Holotype. — USNM 579682, an adult male from Lancetilla (15°44 ′ N, 87°27 ′ W), 30 m elevation, department of Atlántida, Honduras, collected 26 February 2011 by Leonel Marineros. Paratypes (n ¼ 3). — UF 157566, an adult female from Compañia Agricola Paradise near Peña Blanca (14°58 ′ N, 88°01 ′ W), 700 m elevation, Cortés, Honduras; UMMZ 58417, an adult female that is dehydrated and has an incomplete tail, from El Progreso (15°24 ′ N, 87°48 ′ W), Yoro, Honduras, 30 m elevation; MCZ 22045, head and anterior portion of body of an adult, from Progreso District, Yoro, Honduras, about 30 m elevation. Diagnosis. — Tantilla excelsa (in preservative) is defined by the following combination of characters: (1) pale middorsal stripe tan, extending length of body and at least half of tail, stripe on vertebral row and adjacent halves of paravertebral rows throughout its length; (2) tan to white lateral stripe occupying adjacent halves of scale rows 3 and 4; (3) ventrolateral area similar in pale color to that of lateral stripe; (4) lower two-thirds anteriorly and about lower third posteriorly of scale row 1 white similar to that of ventrals; (5) lateral edges of ventrals white, no, or very few, dark spots present; (6) ventral and subcaudal surfaces yellow (shortly after death) and white; (7) two females with 161–178 ventrals, single male with 169 ventrals, subcaudals 61 in one female, 70 in male; (8) tail length 23% of total length in one female, 24% in male. Tantilla excelsa can be distinguished from T. taeniata by having 169 ventrals in single male, 161–178 in two females, and by having yellow ventral surfaces in life (vs. 141–152 ventrals in males and 150 in single known female, and ventral surfaces pale purplish pink grading to darker salmon color on posterior two-thirds of body and tail). Tantilla excelsa differs from T. impensa and T. stenigrammi by having the middorsal pale stripe involving all of the vertebral scale row and adjacent thirds of paravertebral scale rows throughout its length (vs. middorsal stripe confined to vertebral row in those two species [on at least anterior half of body in T. stenigrammi]) and having female ventrals numbering up to 178 (vs. to 172 in T. impensa and 159 in female T. stenigrammi). Tantilla excelsa differs from T. olŋmpia in having complete middorsal and lateral pale stripes, with the middorsal stripe extending onto lateral edges of paravertebral rows, and having 169 ventrals and 70 subcaudal scales in single male (vs. those stripes reduced to dashes and spots, respectively; pale middorsal dashes restricted to vertebral row; and 148 ventrals and 49 subcaudals in a male specimen). Tantilla excelsa differs from T. psittaca by having yellow ventral and subcaudal surfaces in life and white to pale yellow in preservative and in having 169 ventrals in single male and 161–178 in two females (vs. those surfaces pink and grading to red in life and pale brown medially and pale brown with pinkish tinge on posterior third of body and under tail in preservative, 153–163 ventrals in males and 154–161 in female specimens). Tantilla excelsa differs from T. tritaeniata in having 169 ventrals in single male and 161–178 ventrals in two females, and having a complete pale nuchal collar (vs. 157 ventrals in single male and 155–161 in females, and pale nuchal collar divided). Description of holotype. —An adult male; TOL 400 mm; SVL 305 mm; TAL 95 mm (23.8% of TOL); HL 9.8 mm; HW 5.5 mm (level of angle of mouth); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 1.3 mm; snout length 3.0 mm, about 2.3 times longer than eye; pupil circular; rostral about 1.1 times wider than high (2.2 X 2.0 mm); internasal length about 0.8 times width (1.1 X 1.4 mm); prefrontal larger than internasal, wider than long (2.2 X 1.5 mm); median prefrontal suture 1.4 mm, about half as long as frontal; frontal with slight anterior extension, Vshaped posteriorly, about 1.0 times longer than wide (2.9 X 2.8 mm), about 1.0 times longer than distance from its anterior edge to tip of snout (2.9 X 2.8 mm); parietal about 1.5 times longer than wide (4.1 X 2.8 mm), median parietal suture length 3.0 mm, about 1.0 times longer than length of frontal (2.9 mm); parietals contacting five nuchal scales; supraocular about 1.6 times longer than wide (2.3 X 1.4 mm), bordering orbit, contacting parietal, upper postocular, preocular, frontal, and prefrontal. Nasal divided, anterior nasal contacting rostral, internasal, and supralabial 1, posterior nasal contacting internasal, prefrontal, preocular, and supralabials 1 and 2, nostril located in central portion of nasal (forming division of nasal); loreal absent; preocular single, about 1.4 times higher than long (0.7 m X 0.5 mm), lower edge contacting supralabials 2 and 3; postoculars 2, upper about 1.5 times larger than lower (upper height 0.6 mm, lower height 0.4 mm); temporals 1 þ 1, anterior temporal 1.5 times longer than high (1.9 X 1.3 mm), posterior temporal 1.8 times longer than high (1.6 X 0.9 mm); supralabials 7, 3 and 4 bordering orbit, 4 and 5 contacting lower postocular, 5 also contacting anterior temporal, 6 contacting anterior temporal, 7 contacting anterior and posterior temporals; infralabials 6– 7, first 2 contacting anterior pair of chinshields on side with 6 infralabials (left side), first 3 contacting anterior pair of chinshields on side with 7 infralabials (right side); infralabial 1 on each side separated medially by contact with anterior chinshield; mental about 1.6 times wider than long (1.4 X 0.9 mm), contacting first pair of infralabials and anterior pair of chinshields; anterior chinshields about 2.4 times longer than wide (2.6 X 1.1 mm), not extending laterally to border of lip; posterior pair of chinshields about 1.7 times longer than wide (1.7 X 1.0 mm), posterior tips separated from each other by two rows of gular scales; four preventral scales between gular 1 and ventral 1; dorsal scales in 15–15–15 transverse rows, smooth throughout, lacking apical pits and supracloacal tubercles; dorsal scales in six rows at level of subcaudal 10; ventrals 169; cloacal scute divided; subcaudals 70, paired; ventrals plus subcaudals 239. Color of holotype in life. —Other than noting the ventral and subcaudal surfaces were yellow (L. Marineros, personal communication), the collector of the holotype did not record color notes of the recently killed specimen. Color of holotype in preservative (Figs. 6, 7). —Dorsal surfaces brown with white nuchal collar, tan middorsal stripe, and white lateral stripe; tan middorsal stripe includes vertebral row and adjacent tips to a third of paravertebral scale rows; tan middorsal stripe narrowing anteriorly on first scale involved in stripe, extending onto tail, fading out at about midlength of tail; tan middorsal stripe beginning on scale 4 posterior to parietals; broader white lateral stripe present, covering adjacent halves of scale rows 3 and 4, bordered above by continuous, thin dark brown line; white lateral stripe beginning on scale 5 posterior to supralabial 7, extending well onto tail to nearly reaching tip; lower twothirds of scale row 1 white anteriorly, grading to white only reaching lower tips of scale row 1 posteriorly on body; about upper third of scale row 1, all of scale row 2, and lower third of scale row 3 same shade of brown as paravertebral rows on about anterior half of body, before reducing width posteriorly; pale nuchal collar complete, not involving posterior tips of parietals, but involving first two scale rows dorsally, pale nuchal collar also involving posterior half of ultimate supralabial plus two scale rows laterally; dorsal surface of head brown, other than dark brown line crossing posterior tips of parietals to connect with lateral dark head bar; white, with dirty white flecking lateral spot involving postnasal, supralabial 1, and anterior half of supralabial 2; larger white spot posterior to eye involving anterior half of anterior temporal, all of supralabial 5, and tips of supralabials 4 and 6; dark brown lateral bar involving posterior half of supralabial 7, adjacent two-thirds of supralabial 6, and all of posterior temporal to connect with dark brown parietal cross-line; lateral dark bar narrowing only slightly dorsally; anterior tip of mental scale and lateral tips of infralabials 1 and 2 with brown flecking; dark brown spot present on infralabial 4 and adjacent edge of infralabial 5, remainder of ventral surface of head white; ventral surfaces of body and tail white for full length, except for variation discussed above. Variation in paratypes. —Two adult female paratypes show the following variation (followed by mean in parentheses): TOL 326 mm in one (UF 157566; UMMZ 58417 has incomplete tail); SVL 251–276 (263.5). mm; TAL 23.0% of TOL in one; ventrals 161–178 (169.5); subcaudals 61 in one; ventrals plus subcaudals 222 in one. All paratypes (including MCZ R22045, a head and anterior portion of body only of an adult) have 7 supralabials on each side, with supralabials 3 and 4 bordering the eye. Available color pattern notes indicate those paratypes were similar overall to the color pattern of the holotype. Distribution and habitat. —Very little is known about the habitats of Tantilla excelsa. The holotype was killed by a worker at the Lancetilla Botanical Gardens during February. The UF specimen was taken after dark as it was crawling on the ground on a coffee farm during May. The UMMZ specimen was collected on 27 March 1923 and possibly came from an insect trap (G. Schneider, personal communication). No data are available for the MCZ paratype other than it was collected in 1925. The Lancetilla Botanical Gardens formerly contained Lowland Moist Forest and lies at 30 m elevation. The coffee farm is in an area formerly containing Premontane Wet Forest and lies at 700 m elevation. The remaining two localities are in Lowland Dry Forest, although more mesic corridors of gallery forest do occur in those areas. Thus, T. excelsa is known to occur at low and moderate elevations (30–700 m) on both sides of the Río Ulúa drainage, and from the vicinity of Lago de Yojoa, Cortés, to the north-northeast in the vicinity of Lancetilla, Atlántida (Fig. 5). Etymology. —The specific name excelsa is an adjective from the Latin excelsus, which in this case means high. The name refers to the relatively high number of ventral scales in this nominal form., Published as part of McCranie, James R. & Smith, Eric N., 2017, A Review of the Tantilla taeniata Species Group (Reptilia: Squamata: Colubridae: Colubrinae) in Honduras, with the Description of Three New Species, pp. 338-348 in Herpetologica 73 (4) on pages 342-344, DOI: 10.1655/Herpetologica-D-16-00080.1, http://zenodo.org/record/7712317, {"references":["Wilson, L. D., and J. R. Meyer. 1971. A revision of the taeniata group of the colubrid snake genus Tantilla. Herpetologica 27: 11 - 40.","Wilson, L. D., and J. R. Meyer 1982. The snakes of Honduras. Milwaukee Public Museum Contributions in Biology and Geology 6: 1 - 159.","Wilson, L. D., and J. R. Meyer. 1985. The Snakes of Honduras, 2 nd edition. Milwaukee Public Museum, USA.","Wilson, L. D., and J. R. McCranie. 1999. The systematic status of Honduran populations of the Tantilla taeniata group (Serpentes: Colubridae), with notes on other populations. Amphibia-Reptilia 20: 326 - 329.","McCranie, J. R. 2011 a. The Snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, USA.","McCranie, J. R. 2011 b. A new species of Tantilla of the taeniata species group (Reptilia, Squamata, Colubridae, Colubrinae) from northeastern Honduras. Zootaxa 3037: 37 - 44.","Wilson, L. D., and V. Mata-Silva. 2015. A checklist and key to the snakes of the Tantilla clade (Squamata: Colubridae), with comments on taxonomy, distribution, and conservation. Mesoamerican Herpetology 2: 418 - 498."]}

Sigalegalephrŋnus minangkabauensis sp. nov. Holotype. — Museum Zoologicum Bogoriense Amphibian Collection, MZB 25738 (field number ENS 16028), an adult male (Fig. 7) from Gunung Kunyit, Kecamatan Panyabungan Selatan, Kabupaten Kerinci, Provinsi Jambi, Indonesia, 2.26013°S, 101.49512°E at 1402 m, collected by E. Wostl, E.N. Smith, W. Trilaksono, and G. Barraza on 24 June 2013. Diagnosis and comparison. —The following combination of characters is unique to Sigalegalephrŋnus minangkabauensis: (1) A small (19.32 mm SVL) and slender toad without parotoid glands. (2) Fingertips I and II are rounded and not expanded. (3) Fingertips III and IV are rounded and expanded. (4) The toe tips are rounded but not expanded. (5) The webbing is rudimentary in the hands and moderate in the feet. (6) The dorsum is light greenish-brown with a middorsal pinstripe extending from the tip of the snout to the vent. (7) The flanks have a single stroke of dark brown extending from the posterior end of the orbit to the inguinal region. (8) The dorsal surface is moderately tuberculate. (9) The ventral surface is smooth with scattered black spots. Sigalegalephrŋnus minangkabauensis can be distinguished from S. mandailinguensis (characters in parentheses) based on the following differences. The tympanum is barely discernible (tympanum distinct). The finger-pads are moderately defined (finger-pads prominent). The fingertips are rounded but not expanded (distinctly spatulate on tips III to IV). The hands lack subarticular tubercles (distinct subarticular tubercles under Fingers III and IV). The webbing of the foot is more extensive, extending to the last phalanx on Toes I and II (last phalanx free of webbing). The pads on toes are moderately defined (pads prominent). The feet lack subarticular tubercles (distinct subarticular tubercles under Toes IV and V). The overall texture is glossy with fewer tubercles on the dorsum and flanks (overall texture rugose, body and flanks extensively tuberculate). Taking into account the uncorrected genetic distance between the two species of Sigalegalephrŋnus (Table 2), the aforementioned comparisons provide adequate diagnostic characters to warrant S. minangkabauensis as a species distinct from S. mandailinguensis. Description of holotype. —The holotype (sex indistinguishable) has SVL of 19.32 mm; head length 6.94 mm; head width 6.57 mm; snout length 2.7 mm; eye length 2.2 mm; eye–nares length 1.6 mm; distance between nares to tip of snout 0.6 mm; internarial distance 1.8 mm; intercanthal distance 3.6 mm; forearm length 6.0 mm; hand length 5.5 mm; femur length 8.5 mm; tibia length 8.71 mm; tarsal length 5.1 mm; foot length 7.5 mm; width of fingertip pad of Finger III 1.6 mm; Finger IV 1.4 mm. Body slender, head little longer than wide; head length 36.1% SVL, head width 34.0% SVL; snout length 14% SVL; canthus rostralis concave; loreal area without tubercules and concave; snout truncated and slightly sloping back toward mouth; snout mucronate and with prominent median keel in dorsal view; eye length 11.4% SVL; pupil horizontal; upper eyelid granular; tympanum barely visible, with no supratympanic fold; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval-shaped, longer than wide; skin of dorsal surfaces rough to finely shagreen with few large, scattered tubercles; tubercles small, rounded, and almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with few fine warts; forearm length 31.1% SVL; hand length 28.5% SVL; relative lengths of Fingers I Color in life. —Edges of lore and head golden with black shades; area below eyes with prominent white marking with yellowish tint; dorsum light greenish-brown with light brown hourglass figure extending from posterior of orbit to top of the sacroiliac joint; hourglass shape ends with distinct horizontal black bean color mark on each side; yellowishgreen marking on each shoulder; flanks black with red tubercles, maculated with greenish-yellow blotches, and possessing very prominent dark brown stripe starting from posterior end of orbit to inguinal region; inguinal areas greenish with golden tint; sacroiliac joint to inguinal region of flanks, golden yellowish-green; dorsal sides of limbs light brown; forearm, femur, tibia–fibula, and tarsus with distinct dark spot encircled with golden-yellowish-green color; venter opaque; throat golden-yellow; abdomen, ventral side of arms and legs pinkish with scattered yellow and black blotches; webbing in hand and foot translucent. Color in preservative. —In preservative, the animal appears dull because it has lost its golden-yellowish and greenish colors. The hourglass pattern has turned gray. The venter has lost all of its pinkish and golden-yellowish shades and turned a greenish-white. Etymology. —The specific epithet refers to the Minangkabau or Minang ethnic group inhabiting the region where the new species was found. Common name. —Minangkabau Puppet Toads. Province, Sumatra (MZB 25738). A color version of this figure is available online. Distribution and natural history. — Sigalegalephrŋnus minangkabauensis is known only from Gunung Kunyit from an elevation of 1428 m (Fig. 4). The holotype was found perched on a leaf ~ 1.25 m above ground, by the edge of a forest stream at 2015 h. Before collecting it, ENS watched the specimen move in reverse toward the edge of the leaf on which it was perched, where it defecated (outside of the surface of the leaf), and then return to its original position. The holotype weighed 0.5 g., Published as part of Smart, Utpal, Sarker, Goutam C., Arifin, Umilaela, Harvey, Michael B., Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2017, A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a Phylogeny of Sundaland Toad Genera, pp. 63-75 in Herpetologica 73 (1) on pages 63-75, DOI: 10.1655/Herpetologica-D-16-00041, http://zenodo.org/record/7716435

Sigalegalephrŋnus minangkabauensis sp. nov. Holotype. — Museum Zoologicum Bogoriense Amphibian Collection, MZB 25738 (field number ENS 16028), an adult male (Fig. 7) from Gunung Kunyit, Kecamatan Panyabungan Selatan, Kabupaten Kerinci, Provinsi Jambi, Indonesia, 2.26013°S, 101.49512°E at 1402 m, collected by E. Wostl, E.N. Smith, W. Trilaksono, and G. Barraza on 24 June 2013. Diagnosis and comparison. —The following combination of characters is unique to Sigalegalephrŋnus minangkabauensis: (1) A small (19.32 mm SVL) and slender toad without parotoid glands. (2) Fingertips I and II are rounded and not expanded. (3) Fingertips III and IV are rounded and expanded. (4) The toe tips are rounded but not expanded. (5) The webbing is rudimentary in the hands and moderate in the feet. (6) The dorsum is light greenish-brown with a middorsal pinstripe extending from the tip of the snout to the vent. (7) The flanks have a single stroke of dark brown extending from the posterior end of the orbit to the inguinal region. (8) The dorsal surface is moderately tuberculate. (9) The ventral surface is smooth with scattered black spots. Sigalegalephrŋnus minangkabauensis can be distinguished from S. mandailinguensis (characters in parentheses) based on the following differences. The tympanum is barely discernible (tympanum distinct). The finger-pads are moderately defined (finger-pads prominent). The fingertips are rounded but not expanded (distinctly spatulate on tips III to IV). The hands lack subarticular tubercles (distinct subarticular tubercles under Fingers III and IV). The webbing of the foot is more extensive, extending to the last phalanx on Toes I and II (last phalanx free of webbing). The pads on toes are moderately defined (pads prominent). The feet lack subarticular tubercles (distinct subarticular tubercles under Toes IV and V). The overall texture is glossy with fewer tubercles on the dorsum and flanks (overall texture rugose, body and flanks extensively tuberculate). Taking into account the uncorrected genetic distance between the two species of Sigalegalephrŋnus (Table 2), the aforementioned comparisons provide adequate diagnostic characters to warrant S. minangkabauensis as a species distinct from S. mandailinguensis. Description of holotype. —The holotype (sex indistinguishable) has SVL of 19.32 mm; head length 6.94 mm; head width 6.57 mm; snout length 2.7 mm; eye length 2.2 mm; eye–nares length 1.6 mm; distance between nares to tip of snout 0.6 mm; internarial distance 1.8 mm; intercanthal distance 3.6 mm; forearm length 6.0 mm; hand length 5.5 mm; femur length 8.5 mm; tibia length 8.71 mm; tarsal length 5.1 mm; foot length 7.5 mm; width of fingertip pad of Finger III 1.6 mm; Finger IV 1.4 mm. Body slender, head little longer than wide; head length 36.1% SVL, head width 34.0% SVL; snout length 14% SVL; canthus rostralis concave; loreal area without tubercules and concave; snout truncated and slightly sloping back toward mouth; snout mucronate and with prominent median keel in dorsal view; eye length 11.4% SVL; pupil horizontal; upper eyelid granular; tympanum barely visible, with no supratympanic fold; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval-shaped, longer than wide; skin of dorsal surfaces rough to finely shagreen with few large, scattered tubercles; tubercles small, rounded, and almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with few fine warts; forearm length 31.1% SVL; hand length 28.5% SVL; relative lengths of Fingers I Color in life. —Edges of lore and head golden with black shades; area below eyes with prominent white marking with yellowish tint; dorsum light greenish-brown with light brown hourglass figure extending from posterior of orbit to top of the sacroiliac joint; hourglass shape ends with distinct horizontal black bean color mark on each side; yellowishgreen marking on each shoulder; flanks black with red tubercles, maculated with greenish-yellow blotches, and possessing very prominent dark brown stripe starting from posterior end of orbit to inguinal region; inguinal areas greenish with golden tint; sacroiliac joint to inguinal region of flanks, golden yellowish-green; dorsal sides of limbs light brown; forearm, femur, tibia–fibula, and tarsus with distinct dark spot encircled with golden-yellowish-green color; venter opaque; throat golden-yellow; abdomen, ventral side of arms and legs pinkish with scattered yellow and black blotches; webbing in hand and foot translucent. Color in preservative. —In preservative, the animal appears dull because it has lost its golden-yellowish and greenish colors. The hourglass pattern has turned gray. The venter has lost all of its pinkish and golden-yellowish shades and turned a greenish-white. Etymology. —The specific epithet refers to the Minangkabau or Minang ethnic group inhabiting the region where the new species was found. Common name. —Minangkabau Puppet Toads. Province, Sumatra (MZB 25738). A color version of this figure is available online. Distribution and natural history. — Sigalegalephrŋnus minangkabauensis is known only from Gunung Kunyit from an elevation of 1428 m (Fig. 4). The holotype was found perched on a leaf ~ 1.25 m above ground, by the edge of a forest stream at 2015 h. Before collecting it, ENS watched the specimen move in reverse toward the edge of the leaf on which it was perched, where it defecated (outside of the surface of the leaf), and then return to its original position. The holotype weighed 0.5 g.

Sigalegalephrynus gen. nov. Type species. — Sigalegalephrŋnus mandailinguensis by present designation. Diagnosis and comparisons. —The genus can be diagnosed based on the following ecological attributes and morphology: medium-sized (Ansonia (> 40 mm SVL), commonly called Stream toads, is typically found on low vegetation near watercourses. The generally diminutive members of Pelophrŋne (Leptophrŋne (> 40 mm SVL) occur on shrubs. The only other genus of toads that has true arboreal habits is Rentapia, whose members typically tend to be stockier (> 70 mm SVL) than Sigalegalephrŋnus. The genus Phrŋnoidis is represent-ed by two large (> 70 mm SVL) semiaquatic toads, usually found on rocks along streams and rivers; the members of the Duttaphrŋnus (> 40 mm SVL) and Ingerophrŋnus (> 40 mm SVL) display terrestrial or somewhat riparian habits. The monotypic genus Pseudobufo is represented by a large (> 75 mm SVL) and aquatic species with completely webbed feet that inhabits the peat swamps of the Malay Peninsula, Borneo, and Eastern Sumatra. Like all other Sundaland toad genera (characters in parentheses), the new genus possesses a visible or slightly visible tympanum. Sigalegalephrŋnus most closely resembles Ansonia; however, the former lacks mandibular spines (mandibular spines present) and possesses combined femur and tibia lengths smaller than its SVL (SVL Pelophrŋne, in Sigalegalephrŋnus Finger I projects beyond the webbing by two phalanges (reduced Finger I, with one or no phalanges projecting beyond webbing), and males possess nuptial excrescences with well-keratinized spicules (poorly spiculated with only slight keratinization or not keratinized at all). Unlike Rentapia, Sigalegalephrŋnus lacks paratoid glands (paratoids prominent). The new genus can be told apart from Ingerophrŋnus by the lack of well-defined parallel crests between the eyes (parallel crests prominent). Unlike Leptophrŋne, Sigalegalephrŋnus lacks enlarged tubercles at the base of each toe, between at the articulation of the first phalanx and metacarpus (large tubercles present). Additionally, males of L. cruentata are unique among Southeast Asian toads in having nuptial excrescences that are white and swollen on the first and second fingers; males of Sigalegalephrŋnus (and all other genera) have nuptial excrescences with no white and swollen tissue. Unlike Phrŋnoidis, Sigalegalephrŋnus has slender limbs (limbs robust) and toes that are less than half webbed (toes fully webbed, with the exception of the fourth). Unlike Pseudobufo —the only Sundaland toad with fully webbed toes— Sigalegalephrŋnus has toes that are less than half webbed. Males of Sigalegalephrŋnus can be distinguished from all other toads in the region by the presence of an elongate inner metacarpal-thenar tubercle, which is as distinct and large as the outer metacarpal tubercle, and is located medially (Fig. 2). Males of Leptophrŋne cruentata have an elongate and medially located inner metacarpal tubercle, but this is less distinct and noticeably smaller than the outer metacarpal tubercle, whereas Rentapia and Pelophrŋne lack the inner metacarpal tubercle altogether. Fingertips three and four of the new genus are truncated, reflecting arboreality as in Pelophrŋne, Sabahphrŋnus, Rentapia, and some species of Ansonia. Etymology. —The generic name is derived from the name given by the indigenous Batak people of the Toba region in Sumatera Utara to life-sized wooden puppets called Sigale Gale. These puppets are used during the papurpur sepata funerary festivals to placate the spirits of the dead who have left no children behind. The suffix is derived from the masculine and Latinized Greek noun for toad, phrŋnos. The new genus, with a relatively large size compared with most arboreal toads in the region, lanky hands, and a wood-brown complexion, is evocative of the Sigale Gale. Common name. —Puppet Toads. Content. —Our phylogenetic analyses indicate the presence of two species within the new genus: S. mandailinguensis Smart et al.; and S. minangkabauensis Smart et al., Published as part of Smart, Utpal, Sarker, Goutam C., Arifin, Umilaela, Harvey, Michael B., Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2017, A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a Phylogeny of Sundaland Toad Genera, pp. 63-75 in Herpetologica 73 (1) on pages 63-75, DOI: 10.1655/Herpetologica-D-16-00041, http://zenodo.org/record/7716435

Sigalegalephrynus gen. nov. Type species. — Sigalegalephrŋnus mandailinguensis by present designation. Diagnosis and comparisons. —The genus can be diagnosed based on the following ecological attributes and morphology: medium-sized (Ansonia (> 40 mm SVL), commonly called Stream toads, is typically found on low vegetation near watercourses. The generally diminutive members of Pelophrŋne (Leptophrŋne (> 40 mm SVL) occur on shrubs. The only other genus of toads that has true arboreal habits is Rentapia, whose members typically tend to be stockier (> 70 mm SVL) than Sigalegalephrŋnus. The genus Phrŋnoidis is represent-ed by two large (> 70 mm SVL) semiaquatic toads, usually found on rocks along streams and rivers; the members of the Duttaphrŋnus (> 40 mm SVL) and Ingerophrŋnus (> 40 mm SVL) display terrestrial or somewhat riparian habits. The monotypic genus Pseudobufo is represented by a large (> 75 mm SVL) and aquatic species with completely webbed feet that inhabits the peat swamps of the Malay Peninsula, Borneo, and Eastern Sumatra. Like all other Sundaland toad genera (characters in parentheses), the new genus possesses a visible or slightly visible tympanum. Sigalegalephrŋnus most closely resembles Ansonia; however, the former lacks mandibular spines (mandibular spines present) and possesses combined femur and tibia lengths smaller than its SVL (SVL Pelophrŋne, in Sigalegalephrŋnus Finger I projects beyond the webbing by two phalanges (reduced Finger I, with one or no phalanges projecting beyond webbing), and males possess nuptial excrescences with well-keratinized spicules (poorly spiculated with only slight keratinization or not keratinized at all). Unlike Rentapia, Sigalegalephrŋnus lacks paratoid glands (paratoids prominent). The new genus can be told apart from Ingerophrŋnus by the lack of well-defined parallel crests between the eyes (parallel crests prominent). Unlike Leptophrŋne, Sigalegalephrŋnus lacks enlarged tubercles at the base of each toe, between at the articulation of the first phalanx and metacarpus (large tubercles present). Additionally, males of L. cruentata are unique among Southeast Asian toads in having nuptial excrescences that are white and swollen on the first and second fingers; males of Sigalegalephrŋnus (and all other genera) have nuptial excrescences with no white and swollen tissue. Unlike Phrŋnoidis, Sigalegalephrŋnus has slender limbs (limbs robust) and toes that are less than half webbed (toes fully webbed, with the exception of the fourth). Unlike Pseudobufo —the only Sundaland toad with fully webbed toes— Sigalegalephrŋnus has toes that are less than half webbed. Males of Sigalegalephrŋnus can be distinguished from all other toads in the region by the presence of an elongate inner metacarpal-thenar tubercle, which is as distinct and large as the outer metacarpal tubercle, and is located medially (Fig. 2). Males of Leptophrŋne cruentata have an elongate and medially located inner metacarpal tubercle, but this is less distinct and noticeably smaller than the outer metacarpal tubercle, whereas Rentapia and Pelophrŋne lack the inner metacarpal tubercle altogether. Fingertips three and four of the new genus are truncated, reflecting arboreality as in Pelophrŋne, Sabahphrŋnus, Rentapia, and some species of Ansonia. Etymology. —The generic name is derived from the name given by the indigenous Batak people of the Toba region in Sumatera Utara to life-sized wooden puppets called Sigale Gale. These puppets are used during the papurpur sepata funerary festivals to placate the spirits of the dead who have left no children behind. The suffix is derived from the masculine and Latinized Greek noun for toad, phrŋnos. The new genus, with a relatively large size compared with most arboreal toads in the region, lanky hands, and a wood-brown complexion, is evocative of the Sigale Gale. Common name. —Puppet Toads. Content. —Our phylogenetic analyses indicate the presence of two species within the new genus: S. mandailinguensis Smart et al.; and S. minangkabauensis Smart et al.

Sigalegalephrŋnus mandailinguensis sp. nov. Holotype. —Museum Zoologicum Bogoriense Amphibian Collection, MZB 25736 (field number ENS 15697; Fig. 3), an adult male from above the village of Sibanggor Tonga on the northeast slope of Gunung Sorikmarapi, Kecamatan Panyabungan Selatan, Kabupaten Mandailing Natal, Provinsi Sumatera Utara, Indonesia, 0.70164°N, 99.55262°E, 1383 m (in all cases, datum ¼ WGS84), collected by U. Smart, I. Sidik, and E.N. Smith on 25 January 2014. Paratypes (3). — UTA 63561 (field number ENS 16709), adult male from trail between the Tano Bato to Sapo Tinjak road and Lake Saba Begu, Batang Gadis National Park, Kecamatan Batang Natal, Kabupaten Mandailing Natal, Provinsi Sumatera Utara, Sumatra, Indonesia 0.70845°N, 99.51899°E at 1299 m. UTA 63562 (field number ENS 16936) and MZB 25737 (field number ENS 16937), adult males from the same locality as previous paratype but at 1297 m 0.70866°N, 99.51953°E. All paratypes collected by U. Smart, S. Handayani, and I. Sidik on 26 January 2014. Diagnosis. —The following combination of characters is unique to Sigalegalephrŋnus mandailinguensis: (1) This medium-sized (males 30.6–38.0 mm SVL) slender toad lacks parotoid glands. (2) The tympanum is visible. (3) The nares are closer to the tip of the snout than to the eye. (4) The fingertips are truncated and expanded. (5) The toe tips are truncated but not expanded. (6) The webbing is rudimentary in the hands and moderate in the feet. (7) The dorsal coloration consists of white and brown with a thin stripe extending from the tip of the snout to the vent at midline. (8) A dark band above orbits is joined medially, and extends as an interrupted medial track to the sacrum. (9) Alternate black–dark brown and white marks on the upper lip. (10) The flanks have a stroke of brown extending from the orbit to the inguinal area. (11) The dorsal surface is moderately tuberculate. (12) The surface of the abdomen is uniformly tuberculate, with small, smooth, and round tubercles. Description of holotype and variation. — Holotype (adult male) followed by variation of three adult male paratypes in parentheses (UTA 63561; UTA 63562; MZB 25737). The specimen has SVL of 38.0 mm (30.6, 32.29, 32.75); head length 11.34 mm (9.5, 9.73, 10.19); head width 11.3 mm (9.65, 10.27, 10.22); snout length 5.0 mm (4.0, 3.6, 3.5); internarial distance 3.0 mm (2.5, 2.68, 2.75); eye length 3.3 mm (2.8, 3.1, 3); interocular distance 4 mm (3.5, 3.5, 3.7); intercanthal distance 5.7 mm (4.4, 5.0, 5.5); tympanum width 2.1 mm (1.5, 1.8, 1.8); hand length 11.2 mm (9.4, 9.1, 9.8); forearm length 11.5 mm (10, 9.6, 10.3); femur length 15.85 mm (13.46, 12.88, 13.38); tibia length 17.0 mm (13.86, 13.1, 14.02); tarsus length 9.0 mm (7.3, 7.2, 7.5); foot length 16.37 mm (13.58, 13.04, 13.68); width of fingertip pads for Finger I 0.10 mm (0.09, 0.08, 0.09), Finger II 0.13 mm (0.11, 0.10, 0.11), Finger III 0.14 mm (0.11, 0.11, 0.13), Finger IV 0.14 mm (0.10, 0.11, 0.13); width of toe pads for Toe I 0.09 mm (0.07, 0.07, 0.07), Toe II 0.09 mm (0.08, 0.08, 0.08), Toe III 0.11 mm (0.08, 0.08, 0.09), Toe IV 0.12 mm (0.09, 0.09, 0.10), Toe V 0.12 mm (0.09, 0.09, 0.11). Body slender; head almost as long as wide; head length 30% (31%, 30%, 31%) of SVL; head width 30.0% (32%, 32%, 31%) of SVL; snout length 13% (13%, 11%, 11%) of SVL; canthus rostralis concave; loreal area slightly tuberculate and concave; eye length 8.7% of SVL; pupil horizontal; snout slightly sloping back toward mouth; snout mucronate and with prominent median keel, in dorsal view; tympanum distinct, with moderately developed supratympanic fold; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval-shaped and longer than wide; skin of dorsal surfaces rough to finely shagreen with few large, scattered tubercles; tubercles small, rounded, and almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with very fine warts; forearm length 30.3% (32.7%, 29.7%, 31.5%) of SVL; hand length 29.5% (30.7%, 28.2%, 29.9%) of SVL; relative length of Finger I Color in life. —Distinct light brown spot on each side of lore and distinct whitish spot on upper jaw just below posterior end of orbit; dorsum brown with light chamoisbrown-colored hourglass marking; distinct darker spots dorsolateral of sacral and iliac joint; alternate black and brown bands on dorsal side of limbs; in lateral flanks, skin with distinct blackish stripe extending from posterior end of eyes through top of humeral–ulnar joint to inguinal area; short, black dorsal stripe on each side of pubic junction; venter yellowish-white maculated with black blotches; chest and ventral surface of limbs light tea-rose orange with few sparsely located yellowish dots; tips of fingers and toes tearose orange-colored without dots; iris bright gold with black reticulations. Color in preservative. —In preservative the color of the animal differs slightly from that in life. The dorsum has a very light hourglass marking. The venter is whitish grey maculated with dark brown blotches. Etymology. —The specific epithet is an adjective referring to the Mandailing Batak ethnic region and Kabupaten (regency) where the new species was found. Common name. —Mandailing Puppet Toads. Distribution and natural history. — Sigalegalephrŋnus mandailinguensis is known only from rainforests on the slopes of Gunung Sorikmarapi in southern Sumatera Utara from 1297 m to 1383 m (Figs. 4 and 5). The holotype was found at 1830 h inactive on a cave wall, ~ 2 m from the entrance and below ground level. The paratypes were active on vegetation 0.5 m, 2 m, and 3 m (on a liana) above ground between 1930 and 2115 h. The two individuals higher above the ground were vocalizing using a series of high-pitched ticks. The holotype weighed 2.5 g, and the paratypes 1.4, 1.7, and 1.8 g. Call. —The advertisement call of Sigalegalephrynus mandailinguensis is best described as a rapid succession of regularly placed, shrill ticks. A single call 17.27 s in length was recorded at 2156 h. The call begins with infrequent, lowamplitude ticks, which progressively get louder and attain a uniform pulse rate during the main calling period that, in our recording, lasted ~ 15 s. During this time, the tonal pulses were organized in a discreet series repeated at a moderately high rate of 6 times/s. Each individual pulse lasted about 29 ± 4 ms (Fig. 6) intersected by pauses ~ 12 ± 9 ms long. These pulses were relatively narrow band (, Published as part of Smart, Utpal, Sarker, Goutam C., Arifin, Umilaela, Harvey, Michael B., Sidik, Irvan, Hamidy, Amir, Kurniawan, Nia & Smith, Eric N., 2017, A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a Phylogeny of Sundaland Toad Genera, pp. 63-75 in Herpetologica 73 (1) on pages 63-75, DOI: 10.1655/Herpetologica-D-16-00041, http://zenodo.org/record/7716435

Sigalegalephrŋnus mandailinguensis sp. nov. Holotype. —Museum Zoologicum Bogoriense Amphibian Collection, MZB 25736 (field number ENS 15697; Fig. 3), an adult male from above the village of Sibanggor Tonga on the northeast slope of Gunung Sorikmarapi, Kecamatan Panyabungan Selatan, Kabupaten Mandailing Natal, Provinsi Sumatera Utara, Indonesia, 0.70164°N, 99.55262°E, 1383 m (in all cases, datum ¼ WGS84), collected by U. Smart, I. Sidik, and E.N. Smith on 25 January 2014. Paratypes (3). — UTA 63561 (field number ENS 16709), adult male from trail between the Tano Bato to Sapo Tinjak road and Lake Saba Begu, Batang Gadis National Park, Kecamatan Batang Natal, Kabupaten Mandailing Natal, Provinsi Sumatera Utara, Sumatra, Indonesia 0.70845°N, 99.51899°E at 1299 m. UTA 63562 (field number ENS 16936) and MZB 25737 (field number ENS 16937), adult males from the same locality as previous paratype but at 1297 m 0.70866°N, 99.51953°E. All paratypes collected by U. Smart, S. Handayani, and I. Sidik on 26 January 2014. Diagnosis. —The following combination of characters is unique to Sigalegalephrŋnus mandailinguensis: (1) This medium-sized (males 30.6–38.0 mm SVL) slender toad lacks parotoid glands. (2) The tympanum is visible. (3) The nares are closer to the tip of the snout than to the eye. (4) The fingertips are truncated and expanded. (5) The toe tips are truncated but not expanded. (6) The webbing is rudimentary in the hands and moderate in the feet. (7) The dorsal coloration consists of white and brown with a thin stripe extending from the tip of the snout to the vent at midline. (8) A dark band above orbits is joined medially, and extends as an interrupted medial track to the sacrum. (9) Alternate black–dark brown and white marks on the upper lip. (10) The flanks have a stroke of brown extending from the orbit to the inguinal area. (11) The dorsal surface is moderately tuberculate. (12) The surface of the abdomen is uniformly tuberculate, with small, smooth, and round tubercles. Description of holotype and variation. — Holotype (adult male) followed by variation of three adult male paratypes in parentheses (UTA 63561; UTA 63562; MZB 25737). The specimen has SVL of 38.0 mm (30.6, 32.29, 32.75); head length 11.34 mm (9.5, 9.73, 10.19); head width 11.3 mm (9.65, 10.27, 10.22); snout length 5.0 mm (4.0, 3.6, 3.5); internarial distance 3.0 mm (2.5, 2.68, 2.75); eye length 3.3 mm (2.8, 3.1, 3); interocular distance 4 mm (3.5, 3.5, 3.7); intercanthal distance 5.7 mm (4.4, 5.0, 5.5); tympanum width 2.1 mm (1.5, 1.8, 1.8); hand length 11.2 mm (9.4, 9.1, 9.8); forearm length 11.5 mm (10, 9.6, 10.3); femur length 15.85 mm (13.46, 12.88, 13.38); tibia length 17.0 mm (13.86, 13.1, 14.02); tarsus length 9.0 mm (7.3, 7.2, 7.5); foot length 16.37 mm (13.58, 13.04, 13.68); width of fingertip pads for Finger I 0.10 mm (0.09, 0.08, 0.09), Finger II 0.13 mm (0.11, 0.10, 0.11), Finger III 0.14 mm (0.11, 0.11, 0.13), Finger IV 0.14 mm (0.10, 0.11, 0.13); width of toe pads for Toe I 0.09 mm (0.07, 0.07, 0.07), Toe II 0.09 mm (0.08, 0.08, 0.08), Toe III 0.11 mm (0.08, 0.08, 0.09), Toe IV 0.12 mm (0.09, 0.09, 0.10), Toe V 0.12 mm (0.09, 0.09, 0.11). Body slender; head almost as long as wide; head length 30% (31%, 30%, 31%) of SVL; head width 30.0% (32%, 32%, 31%) of SVL; snout length 13% (13%, 11%, 11%) of SVL; canthus rostralis concave; loreal area slightly tuberculate and concave; eye length 8.7% of SVL; pupil horizontal; snout slightly sloping back toward mouth; snout mucronate and with prominent median keel, in dorsal view; tympanum distinct, with moderately developed supratympanic fold; interorbital space flat; cranial crests absent; no teeth in jaws; tongue tip oval-shaped and longer than wide; skin of dorsal surfaces rough to finely shagreen with few large, scattered tubercles; tubercles small, rounded, and almost without keratinization; no dorsolateral, paravertebral, or occipital folds; skin on venter smooth with very fine warts; forearm length 30.3% (32.7%, 29.7%, 31.5%) of SVL; hand length 29.5% (30.7%, 28.2%, 29.9%) of SVL; relative length of Finger I Color in life. —Distinct light brown spot on each side of lore and distinct whitish spot on upper jaw just below posterior end of orbit; dorsum brown with light chamoisbrown-colored hourglass marking; distinct darker spots dorsolateral of sacral and iliac joint; alternate black and brown bands on dorsal side of limbs; in lateral flanks, skin with distinct blackish stripe extending from posterior end of eyes through top of humeral–ulnar joint to inguinal area; short, black dorsal stripe on each side of pubic junction; venter yellowish-white maculated with black blotches; chest and ventral surface of limbs light tea-rose orange with few sparsely located yellowish dots; tips of fingers and toes tearose orange-colored without dots; iris bright gold with black reticulations. Color in preservative. —In preservative the color of the animal differs slightly from that in life. The dorsum has a very light hourglass marking. The venter is whitish grey maculated with dark brown blotches. Etymology. —The specific epithet is an adjective referring to the Mandailing Batak ethnic region and Kabupaten (regency) where the new species was found. Common name. —Mandailing Puppet Toads. Distribution and natural history. — Sigalegalephrŋnus mandailinguensis is known only from rainforests on the slopes of Gunung Sorikmarapi in southern Sumatera Utara from 1297 m to 1383 m (Figs. 4 and 5). The holotype was found at 1830 h inactive on a cave wall, ~ 2 m from the entrance and below ground level. The paratypes were active on vegetation 0.5 m, 2 m, and 3 m (on a liana) above ground between 1930 and 2115 h. The two individuals higher above the ground were vocalizing using a series of high-pitched ticks. The holotype weighed 2.5 g, and the paratypes 1.4, 1.7, and 1.8 g. Call. —The advertisement call of Sigalegalephrynus mandailinguensis is best described as a rapid succession of regularly placed, shrill ticks. A single call 17.27 s in length was recorded at 2156 h. The call begins with infrequent, lowamplitude ticks, which progressively get louder and attain a uniform pulse rate during the main calling period that, in our recording, lasted ~ 15 s. During this time, the tonal pulses were organized in a discreet series repeated at a moderately high rate of 6 times/s. Each individual pulse lasted about 29 ± 4 ms (Fig. 6) intersected by pauses ~ 12 ± 9 ms long. These pulses were relatively narrow band (

Harvey, Michael B., O'Connell, Kyle A., Wostl, Elijah, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N., Grismer, L. Lee (2016): Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and Phylogeny of the Prehensile-tailed Cyrtodactylus. Zootaxa 4107 (4): 517-540, DOI: http://doi.org/10.11646/zootaxa.4107.4.3

Cyrtodactylus lateralis (Werner) Gymnodactylus lateralis Werner 1896: 11. Holotype female (ZMB 12029, Fig. 1, 2, examined) from ���Sumatra.��� [herein traced to Binjai, Sumatera Utara, Indonesia]. Gymnodactylus lateralis Werner: Werner 1900: 499; Barbour 1912: 80; De Rooij 1915: 7. Cyrtodactylus lateralis (Werner): Kluge 1993: 8; Manthey and Grossmann 1997: 223; R��sler 2000: 66; Teyni�� et al. 2010: 33. Referred specimens examined (19). Seven males (MZB 13171, MZB 13173; UTA 62916, UTA 62919 ���62921) and six females (MZB 13172, MZB 13174 ���13176, MZB 18330, UTA 62917 ���62918) from trail up SE slope of Gunung Seulawah Agam, Aceh Besar Regency, Aceh Province, Sumatra, Indonesia, 685���1065 m collected 30 July and 1 August 2015 by M. B. Harvey and E. Wostl and one female (UTA 62915) from Gunung Batee Meucica, Aceh Besar Regency, Aceh Province, Sumatra, Indonesia, 5.26157 �� N, 95.54176 �� E, 498 m collected 1 August 2015 by G. C. Sarker and I. Sidik; two males (ZMB 65273 ���65274) and two females (ZMB 65271 ���65272) from Gurah, Aceh Province, Sumatra, Indonesia, collected January 1997 by C. Sch��fer and W. Grossmann (traced to a village of the same name in Kecamatan Peukan Bada, Kabupaten Aceh Besar). One female (ZMB 49125) from Ketambe, Aceh Province, Sumatra, Indonesia, collected March 1990 by W. Denzer, C. Steiof and U. Manthey. Diagnosis. A large species of Cyrtodactylus reaching 100 mm SVL (LSUHC 12579) and distinguished from all congeners by the following combination of characters: (1) body robust; limbs and digits moderate in length; (2) tail prehensile; intact tails 0.97���1.22 times longer than body; (3) tubercles extending from frontal region and supraorbital skin to cover most of tail, 16���20 irregular longitudinal rows of tubercles at midbody, 21���28 irregular transverse rows between limbs; (4) dorsal antebrachium and brachium tuberculate; (5) tubercles on tail extending to 90 % of its length; (6) 51���66 ventrals in a transverse row at midbody; (7) conical, spinose tubercles in ventrolateral fold; (8) subcaudals not transversely enlarged; (9) moderate longitudinal sulcus in precloacal region of males; (10) 9���13 precloacal pores in males (0���15) pore primordia in females), femoral pores absent, precloacal pores sunken into precloacal sulcus; pore secretions red; (11) greatly enlarged precloacal pore-bearing scales present; (12) ventral surface of thighs entirely granular or granular except for 2���13 (count combined for both sides) widely spaced enlarged femoral scales; (13) 18���24 lamellae under fourth toe; (14) cloacal tubercles 1���2 on each side, usually in contact with one another; (15) dorsum gray to brown with dark brown markings; venter pale pink, immaculate or with diffuse, darkly pigmented scales forming no obvious pattern; (16) labials pale, contrasting with darker facial band; occiput with few large blotches; postocular stripe brown, edged dorsally by thin black line then by pale pigmented scales; (17) 5���6 transverse, usually broken bands on body; 7���9 bands on tail; caudal bands complete ventrally though faint. Comparisons. Cyrtodactylus lateralis is very similar to C. durio. Cyrtodactylus lateralis has rounded to subpyramidal tubercles in the temporal region, on the occiput, and on the neck, whereas tubercles in these areas are noticeably larger and distinctly spinose in C. durio. We scored three new specimens of C. durio for the same characters used in the description of C. lateralis. Ranges of meristic and mensural characters of C. durio overlap those of C. lateralis, however extremes of three characters were slightly higher or lower than those of C. lateralis. The three specimens of C. durio (data for KU 173086 followed by KU 173087 then LSUHC 12579 vs. ranges [mean �� standard deviation, sample size] for C. lateralis) had slightly fewer tubercles at midbody (17, 15, and 15 vs 16���20 [18 �� 1, n = 13]), slightly fewer lamellae under finger IV (15, 17, and 17 vs. 16���20 [18 �� 1, n = 14]), and a slightly larger orbit (diameter of orbit 27.8 %, 28.7 %, and 26.5 % of head length vs. 20.6���27.6 % [25.4 �� 1.6, n = 16]) than the series of C. lateralis. All specimens of C. durio lack enlarged subfemoral scales, whereas 70 % of C. lateralis had these scales. Cyrtodactylus lateralis, C. brevipalmatus, C. durio, C. elok, and C. stresemanni are the only Sunda Shelf species with a spinose, prehensile tail. Unlike C. stresemanni (characters in parentheses), C. lateralis has 16���20 tubercles across midbody (13), a transversely banded dorsum (wide, dark brown chevrons on flanks separated by undulating pale vertebral stripe), and small irregular blotches on the occiput (large, dark triangular occipital mark). Unlike C. brevipalmatus and C. elok, C. lateralis has a precloacal sulcus (absent), spinose tubercles in the ventrolateral fold (tubercles absent from fold), 51���66 ventrals (35���49), and caudal whorls of six tubercles separated by granular scales ventrolaterally (caudal whorls of four tubercles; ventrolateral tubercles separated by lanceolate scales forming ventrolateral fringe). Description. Characteristics of the holotype placed in brackets follow those of all specimens examined. Large Cyrtodactylus reaching at least 93 mm SVL in females [77 mm], 82 mm in males; smallest specimen subadult female, 62.1 mm SVL; body robust, accounting for 38���49 % (45.6 �� 3.7, n = 7) of SVL in males, 43.0��� 53.9 % (48.2 �� 3.8, n = 8) [49.4 %] of SVL in females; tail 0.99���1.14 (n = 8) [0.97] times longer than body and accounting for 49.3���55.1 % (51.8 �� 2.3, n = 8) [49.3 %] of total length; head triangular in dorsal view, much wider than body and distinct from neck; head 28.9 ���34.0% (30.8 �� 1.7, n = 7) of SVL, 63.9���78.9 % (70.2 �� 6.1, n = 7) as wide as long, and 36.2���43.2 % (38.8 �� 2.6 n = 7) as deep as wide in males and 28.5���33.4 % (31.0 �� 1.7, n = 9) [30.1 %] of SVL, 64.3���71.6 % (69.4 �� 2.7, n = 9) [64.3 %] as wide as long, and 38.0��� 46.2 % (41.6 �� 2.8, n = 9) [41.1 %] as deep as wide in females. Rostral wider than or about as wide as mental (96.8���126.81 %, 106.8 �� 8.5, n = 17, of width of mental) [103.2 %], posteriorly forming border of nostril; dorsal margin of rostral notched medially by shallow groove; rostral groove bifurcating slightly ventral to center of nostril; rostral bordered posteriorly by four (50 %, n = 16), five (31 %), six (6 %), or seven (13 %) postrostrals [rostral irregular in holotype; rostral partially fragmented into two large plates just anterior to straight row of 5 postrostrals; short, bifurcating groove below these large irregular plates; rostral of ZMB 49125 also irregular, postrostral series fused to rostral, large diamond-shaped plate partially fractured within rostral in front of fused postrostrals]; internarial region narrow, 10.4���18.8 % (15.7 �� 2.0, n = 17) [17.6 %] as wide as head, 55.6���70.3 % (63.8 �� 4.5, n = 17) [60.9 %] as wide as narrowest point between orbits; narrowest point between orbits 18.7���28.9 % (24.5 �� 2.7, n = 17) [28.9 %] as wide as head; deep depression occupying most of prefrontal and frontal areas; prefrontal-frontal depression bordered laterally by raised canthal and circumorbital ridges, bordered posteriorly by slightly arched, transverse ridge located just behind orbits; central parietal region slightly depressed; scales on dorsal snout granular, rounded, smooth, subequal in size, about as large as tubercles in parietal regions and distinctly smaller than granules between eyes; tubercles extending to frontal region and supraorbital skin; tubercles on dorsal surface of head subpyrimidal and smooth, much smaller than those on body, those in occipital region surrounded by seven or eight granular scales; few tubercles around back of head subconical; dorsal body tubercles unicarinate, circular, slightly projecting from dorsum, about ten times as large as dorsal granules, 16���20 (18 �� 1, n = 13) [18] longitudinal rows at midbody including tubercles in ventrolateral folds, 21���28 (23 �� 2, n = 13) [27] irregular transverse rows between limbs; dorsal tubercles on flanks like those middorsally or attenuated into conical tubercles; dorsal granules smooth. Nostril oval, directed postero-laterally and somewhat dorsally; bordered by rostral, postrostral, supranasal, postnasals, and narrow antero-dorsal process of first supralabial; supranasal rounded to rhomboidal, about four times as large as adjacent postnasals; postnasals granular, 5���7 (6 �� 1, n = 19) [5 / 5]; snout length 36.7���41.2 % (39.3 �� 1.4, n = 16)[39.4 %] of head length and distance from orbit to nostril accounting for 71.2���78.4 % (74.1 �� 2.0, n = 17) [75.3 %] of snout length; loreal region sloping, concave just behind nostril; lips flared; loreals granular, juxtaposed; 19���25 (21.9 �� 2, n = 21) [21 / 21] loreals between nasal and anterior border of orbit; lorilabials in front of orbit flat, plate-like, enlarged relative to other loreals (1.5���2 times as large as loreals above them, but much smaller than supralabials); eye separated from supralabials by one or two longitudinal rows of tiny granules; diameter of orbit 20.6���27.6 % (25.4 �� 1.6, n = 16) [26.3 %] of head length; temporal region with granular scales separating rounded to subpyrimidal, smooth or weakly keeled tubercles; distance from posterior border of orbit to anterior border of auditory meatus 23.6���31.3 % (29.0 �� 1.9, n = 16) [30.9 %] of head length; skin fold overlapping dorsal edge of auditory meatus; auditory meatus elliptical, it���s greatest diameter vertical or directed posterodorsally less than 20 �� to vertical, positioned at posterior end of head and with its dorsal margin at same level as rictus; greatest diameter of auditory meatus 17.7���39.6 % (25.6 �� 5.8, n = 17) [25.5 %] of diameter of orbit; smallest diameter of auditory meatus 60���70 % smaller than greatest diameter; external ear canal deep; supralabials 8���12 (10 �� 1, n = 21) [9 / 10] to center of orbit; total enlarged supralabials 10���14 (12 �� 1, n = 21) [11 / 12], last separated by 2���5 small granular scales from rictus (13���17 total supralabials to rictus) [15 / 16]; first supralabial with anterodorsal extension reaching nostril and separating lorilabials from rostral; remaining supralabials subrectangular with rounded dorsal edges, reducing in size to anterior border of orbit where one or two supralabials somewhat longer than rest. Mental triangular; infralabials 7���9 to center of orbit (8 �� 1, n = 22) [7 / 7], 9���13 (10 �� 1, n = 22) [10 / 10] total enlarged infralabials to rictus; first pair of postmentals in contact medially, length of their contact about half as long as mental; second pair of postmentals as long as first pair but extending medially only one-half to two-thirds as far, separated from one another medially by 2���7 (6 �� 1, n = 17) [5] small granular scales bordering first pair of postmentals; sublabial row decreasing in size posteriorly, not differentiated from adjacent gulars at level of eye; gulars small, smooth, granular, nonimbricate, much smaller than ventrals; in front of pectoral girdle, gulars grading into progressively larger, imbricate scales; scale of chest like ventrals; ventrals smooth, imbricate, flat, circular, about three-fourths as wide as tubercles on dorsum at midbody; ventrals in transverse row at midbody 51���66 (55 �� 3) [59]; ventrolateral skin fold distinct, containing long spinose tubercles separated by granular scales intermixed with smaller spinose tubercles, subpyrimidal tubercles, and smooth rounded tubercles (Fig. 2); tubercles of ventrolateral fold lacking keels. Tail cylindrical, though somewhat flattened dorsally and ventrally, with distinct lateral sulcus proximally for about two-thirds its length; caudal tubercles 2���3 times as large as those on body, heavily keeled, and projecting, separated from one another by granular scales; caudal tubercles on about 90 % of intact tails, disposed in six longitudinal rows and in regular whorls, one tubercle of each whorl below lateral sulcus on each side; 2���3 tubercles in irregular transverse row between successive whorls on proximal one-third to one-half of tail; tubercles between whorls same shape but only one-half to one-third as large as tubercles forming whorls; size and distribution of tubercles at base of tail like posterior body; subcaudals smooth, flat, homogenous, imbricate, 2���3 times as large as granules on sides of tail; each subcaudal about as long as wide; transversely enlarged subcaudal plates absent; cloacal tubercles rounded, smooth, one (7 %, n = 27) or two (93 %) on each side, contacting one another (80 %, n = 25 paired tubercles) or separated from one another by a single granular scale (20 %) [2 / 2 cloacal tubercles contacting one another on each side of holotype]. Tubercles of limbs keeled, subpyramidal, largest same size as those on dorsum; tuberculation of brachium variable, almost entirely granular to as tuberculate as antebrachium [brachium of holotype tuberculate, but not as heavily tuberculate as antebrachium]; antebrachium tuberculate; antebrachial tubercles about six times as large as granular scales between them; granules and tubercles grading to dorsal patch of about seven rows of large juxtaposed scales covering distal antebrachium and proximal hand; ventral surfaces of brachium covered in smooth, slightly imbricate, homogenous granules; tubercles extending across postaxial and ventral surfaces of antebrachium to approximate position of radius, then transitioning to granular scales on preaxial side of antebrachium; scales on palm juxtaposed to slightly imbricate; lamellae under finger I 8���9 [8], under finger II 12���16 [13], under finger III 15���19 [16], under finger IV 16���20 [17], under finger V 13���16 [14]; one or two scales widened and similar to lamellae extending onto palm at base of digits. Thigh, shank, and postaxial half of dorsal foot (at base of toes IV���V) tuberculate; tubercles of leg subpyrimidal, unicarinate, largest about same size as those on dorsum, about six times as large as small granular scales between them; tubercles on thigh decreasing in size postaxially, grading to tiny granular scales on postaxial edge of thigh (no sharp transition along postaxial edge of thigh); ventral thigh entirely granular (30 %, n = 23) or granular except for row of 1���7 (3���13 total count including both sides; 70 % of the time; Fig. 3) flat enlarged subfemoral scales [ventral thigh entirely granular in holotype, lacking enlarged subfemoral scales]; enlarged subfemoral scales separated from one another by gaps occupied by granular scales; gaps between subfemorals increasing in width distally; sole covered in flat, juxtaposed to weakly imbricate, rounded scales; lamellae under toe I 8���10 [10], under toe II 11���16 [13], under toe III 16���19 [16]; under toe IV 18���24 [19], under toe V 17���20 [18]; one or two scales widened and similar to lamellae extending onto sole at base of toes II���V, three or four extending onto sole at base of toe I. Scales of precloacal region forming patch (Fig. 2, 3) of enlarged, smooth, imbricate scales grading to smaller ventrals anteriorly and small imbricate scales edging vent posteriorly; males with parallel series of 9���13 total precloacal pores; pore-bearing series of males sunken into shallow longitudinal sulcus; same scales of females with 0���15 pore primordia lacking secretions [no detectable pore or pore primordia in holotype]; females lacking longitudinal sulcus; scales flanking pore-bearing scales smaller than pore-bearing scales anteriorly, subequal to them posteriorly; both sexes with pair of post-cloacal slits; males with post-cloacal ossicle in wall of each slit. Coloration. In life (based on field notes of M.B. Harvey and photos of all specimens collected by the summer 2015 expedition to Aceh), Cyrtodactylus lateralis resembles tree bark. This species has a gray, light brown, or reddish brown ground coloration overlain by diffuse, fine darker speckling in addition to bold, dark brown blotches and pale areas largely devoid of speckling (Fig. 4). Dorsal tubercles are the same color as adjacent scales. Dorsally, a few, small, dark brown blotches mark the parietal and occipital regions; however, the rest of the dorsal head is usually devoid of solid dark brown blotches [holotype faded to cream with light brown markings]. The supraciliaries are suffused with orange; they may be almost entirely orange in young specimens, whereas brown pigment obscures the orange supraciliary pigment in older individuals. A wide band extends from the rostral to the eye and continues as a postocular stripe to the arm. The facial band is always weakly defined and may be quite diffuse in some specimens [loreal region of holotype brownish]. Behind the eye, weak, thin, and often broken black pinstripes edge the postocular band dorsally and ventrally [postocular band with black pinstripes extending above and slightly behind arm]. Several rows of temporals above the black pinstripe are cream, pale gray, or pale yellow. These pale temporals form a band behind the eye that breaks up into subcircular spots at the back of the head and on the neck. The labials are also pale, almost immaculate to heavily mottled but always contrasting with the facial band and gular skin [bands of melanophores covering supralabials 5���6, anterior portion of 7, and 8���9 on left/ 1���2 and 9���10 on right; chin diffusely speckled with melanophores]. In specimens with mottled labials, subcircular cream, pale gray, or pale yellow spots, each about as large as a supralabial, mark the labial series. The iris is grayish tan or tan, heavily reticulated with black lines. The ciliary body is only slightly lighter than the iris and indistinct. The tongue is white and lacks the black ventral patch described for some congeners (Harvey et al. 2015); the buccal epithelium is pink. Cyrtodactylus lateralis has a broken V-shaped mark on the nape, one narrow band on the neck, 5���6 [5] bands on the body, and 7���9 (8 �� 1) [7] bands on the tail. Bands on the neck and body usually break up into oval blotches, and pale areas separate the bands middorsally and on the upper flanks. Bands on the body usually have a ���butterflylike��� shape [Fig. 1, 4, see also plate 1, figure 4 of Werner, 1896]. The lower flanks are darkly pigmented and frequently sharply contrast with a pale ventrolateral fold or a fold of the same color with pale yellow or cream spinose tubercles [fold of holotype like flanks, covered in melanophores but with large spinose tubercles of fold white and lacking melanophores]. Caudal bands are square to rectangular, narrower than the pale intersp, Published as part of Harvey, Michael B., O'Connell, Kyle A., Wostl, Elijah, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N. & Grismer, L. Lee, 2016, Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and Phylogeny of the Prehensile-tailed Cyrtodactylus, pp. 517-540 in Zootaxa 4107 (4) on pages 520-527, DOI: 10.11646/zootaxa.4107.4.3, http://zenodo.org/record/256578, {"references":["Werner, F. (1896) Zweiter Beitrag zur Herpetologie der indo-orientalischen Region. Verhandlungen der Zoologische- Botanischen Gesellschaft in Wien, 46, 6 - 24. http: // dx. doi. org / 10.5962 / bhl. part. 19752","Werner, F. (1900) Reptilien und Batrachier aus Sumatra. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere, 13, 479 - 508.","Barbour, T. (1912) A contribution to the zoogeography of the East Indian islands. Memoirs of the Museum of Comparative Zoology, 44, 1 - 203, pls. 1 - 8.","De Rooij, N. de (1915) The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill, Leiden, xiv + 384 pp.","Kluge, A. G. (1993) Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.","Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Sudostasiens. Natur und Tier Verlag, Munster, 512 pp.","Rosler, H. (2000) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28 - 153.","Teynie, A., David, P. & Ohler, A. (2010) Notes on a collection of amphibians and reptiles from Western Sumatra (Indonesia), with the description of a new species of the genus Bufo. Zootaxa, 2416, 1 - 43.","Harvey, M. B., O'Connell, K. A., Barraza, G., Riyanto, A., Kurniawan, N. & Smith, E. N. (2015) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny. Zootaxa, 4020, 495 - 516. http: // dx. doi. org / 10.11646 / zootaxa. 4020.3.5","Cameron, P. (1906) A new genus of Cryptinae (Ichneumonidae) from Sumatra. The Entomologist, 39, 196 - 197. http: // dx. doi. org / 10.5962 / bhl. part. 1560","Grismer, L. L. (2008) On the distribution and identification of Cyrtodactylus brevipalmatus Smith, 1923 and Cyrtodactylus elok Dring, 1979. The Raffles Bulletin of Zoology, 65, 177 - 179.","Grismer, L. L. (2011) Lizards of Peninsular Malaysia, Singapore and their Adjacent Archipelagos. Edition Chimaira, Frankfurt, 728 pp.","Grismer, L. L., Anuar, S., Quah, E., Muin, M. A., Onn, C. K., Grismer, J. L. & Ahmad, N. (2010) A new spiny, prehensile-tailed species of Cyrtodactylus (Squamata: Gekkonidae) from Peninsular Malaysia with a preliminary hypothesis of relationships based on morphology. Zootaxa, 2625, 40 - 52.","Linkem, C. W., McGuire, J. A., Hayden, C. J., Mohammed, I. S., Bickford, D. P. & Brown, R. M. (2008) A new species of bent-toe gecko (Gekkonidae: Cyrtodactylus) from Sulawesi Island, eastern Indonesia. Herpetologica, 64, 224 - 234. http: // dx. doi. org / 10.1655 / 07 - 040.1"]}

Phylogeny of Cyrtodactylus lateralis and other Prehensile-tailed Cyrtodactylus Currently, the Reptile Database lists 206 species of Cyrtodactylus (Uetz & Ho��ek 2015). Systematists face considerable challenges when trying to identify derived morphological characters in this diverse gekkonid radiation. On the basis of morphology, Grismer et al. (2010) proposed a phylogeny of Cyrtodactylus with spinose prehensile tails. These authors constructed their tree using parsimony and polarized a small matrix of characters by treating all other congeners as outgroups. In their phylogeny, Cyrtodactylus brevipalmatus and C. elok form a clade and share digital webbing, a square tail, and a caudal fringe. Cyrtodactylus spinosus, C. stresemanni, and C. durio form a second clade and share spinose tubercles in the ventrolateral fold on the body. Within the second clade, Cyrtodactylus durio and C. stresemanni are sister species and share enlarged, ventrolateral caudal tubercles. The authors argued that a prehensile tail is a synapomorphy of these five species. Using their methods (Table 1), we add Cyrtodactylus lateralis (Werner), C. interdigitalis Ulber, C. nuaulu Oliver et al., and C. serratus Kraus to this morphological phylogeny (Fig. 6). Ulber (1993) compared C. interdigitalis to C. brevipalmatus and C. elok and noted that, unlike these two species, C. interdigitalis has a round rather than square tail. All three species share digital webbing and a ventrolateral fringe. The ventrolateral fringe consists of a distinctive row of enlarged conical tubercles separated by smaller lanceolate tubercles. The enlarged tubercles are the ventrolateral components of caudal whorls, whereas the smaller conical tubercles are the only enlarged scales separating the whorls dorsally. All three species lack spinose tubercles in the ventrolateral fold on the body. Ulber (1993) did not mention whether C. interdigitalis coils its tail at rest, however we have seen photos of specimens from Laos exhibiting this behavior. Based on this combination of traits, we add C. interdigitalis to the morphological phylogeny of Grismer et al. (2010) and place C. interdigitalis as the sister species to a clade containing C. elok and C. brevipalmatus (Fig. 6). We refer to this group of three species as the ���mainland SE Asian clade��� of prehensile-tailed Cyrtodactylus, because these species occur in Laos, Myanmar, Peninsular Malaysia, and Thailand. In their phylogeny, Grismer et al. (2010) placed Cyrtodactylus durio as the sister species to C. stresemanni with C. spinosus sister to the C. durio + C. stresemanni clade. They note that C. spinosus lacks ventrolateral caudal tubercles, but did not explain how they scored this character. We count tubercles in the middle of the tail: C. lateralis, C. durio, and C. stresemanni have six spinose tubercles per caudal whorl throughout most of the tail (the count will be higher close to the body and lower distally), whereas C. spinosus has only 4 (also noted by Linkem et al. 2008). We add C. lateralis to the C. durio + C. stresemanni Clade, because C. durio and C. lateralis are identical for the characters used to construct the morphological phylogeny (Table 1). Cyrtodactylus nuaulu and C. serratus, both from east of Wallace���s Line, have conical tubercles in the ventrolateral fold, ventrolateral caudal tubercles, and tails with whorls of six spinose tubercles extending for most of the length of the tail. On the basis of morphology, we infer that the tails of these two species are prehensile, even though neither description mentions whether the species coils its tail in a manner similar to C. spinosus and the Sunda Shelf species. The unique holotype of C. serratus was collected in the Star Mountains of New Guinea in 1987 and Kraus (2007) provided no information about its behavior or natural history. All adults of the type series of C. nuaulu were collected on trees and vines, 50���200 cm aboveground on Seram (Oliver et al. 2009). Based on these observations, we add C. nuaulu and C. serratus to the clade containing C. stresemanni, C. durio, and C. lateralis. We refer to this clade as the ���insular clade��� of prehensile-tailed geckos, because most of the species occur on islands. Of the five species of the insular clade, only C. durio occurs on the mainland. We have shown that some distinctive characters originally suggesting a close relationship between Cyrtodactylus durio and C. stresemanni are also found in C. serratus and C. nuaulu. Although the specimen has faded with age, the dorsal pattern of the holotype of C. stresemanni remains well-defined (R��sler & Glaw 2008, their figure 7). The specimen has two pairs of elongate blotches on the flanks separated by a wide, pale vertebral stripe. This dorsal pattern is unlike that of any other Sunda Shelf Cyrtodactylus, however it resembles patterns in the type series of C. nuaulu (compare especially to MZB 2325 in Oliver et al. 2009, their figure 2), C. spinosus (see especially MZB 7029 with ���less complete��� chevrons, Linkem et al. 2008: 228), and C. serratus (Kraus 2007, his figure 1 A). All four species share a wide pale vertebral stripe and few (2���3 between limbs) large, dark-brown chevron-shaped blotches. The vertebral stripe may be interrupted by the chevrons medially or divide the chevrons as it does in the holotypes of C. serratus and C. stresemanni and some paratypes of C. spinosus (MZB 7029) and C. nuaulu (MZB 2329). Although the dark triangular blotch on the nape of C. stresemanni might at first appear to ally it with C. elok and C. brevipalmatus, C. nuaulu, C. serratus, and C. spinosus have triangular blotches on their napes as well. Cyrtodactylus durio and C. lateralis unequivocally lack the distinctive color pattern of C. stresemanni and the three other species. Like C. brevipalmatus, C. elok, and C. interdigitalis, C. durio and C. lateralis are banded species lacking a pale vertebral stripe. To determine polarity of the distinctive color pattern of the Wallacean and Papuan species, we reviewed recent descriptions and revisions of Cyrtodactylus from across the distribution of the genus. We were unable to find other species with a pale vertebral stripe and 2���3 dark brown chevrons. C. zugi Oliver has two large brown chevrons, but lacks a vertebral stripe. The pattern of the Wallacean and Papuan prehensile-tailed Cyrtodactylus is most likely derived, however, its distribution is not congruent with the morphological phylogeny regardless of polarity. If derived, the pattern is a synapomorphy of the insular clade, with a reversal to the banded pattern in C. durio and C. lateralis. If the pattern is primitive, a banded pattern would have to evolve twice: once leading to the mainland SE Asia clade and once leading to a clade containing C. durio and C. lateralis. Prior to this study, molecular data were available for only two of the eight species of prehensile-tailed Cyrtodactylus. Wood et al. (2012) confirmed the suspicions of Ulber (1993) that C. elok is closely related to C. interdigitalis, and these two species are sister taxa in Clade F of their phylogeny. We obtained ND 2 and COI sequences for four of our new specimens of C. lateralis and an ND 2 sequence of the holotype of C. durio. Our maximum likelihood and Bayesian analyses recovered similar topologies with minor differences in support for each gene. Relatively few sequences of the COI gene were available on GenBank (Appendix 1), however we were able to place Cyrtodactylus elok and C. lateralis within a phylogeny containing 34 additional congeners and Gekko gekko as an outgroup. Our Maximum Likelihood tree contained few mainland species and had low support for most nodes. Nonetheless, we found no evidence of a close relationship between C. elok and C. lateralis using the COI sequences. In this small matrix of species, C. elok and C. lateralis were placed near the base of the phylogeny and had uncorrected genetic distances greater than 19 % to other species in the phylogeny (Table 2). Our ND 2 matrix included Cyrtodactylus durio, C. elok, C. interdigitalis, C. lateralis, 83 additional congeners, 26 gecko species as near outgroups, and Anolis carolinensis as a distant outgroup. We only discuss the region of the tree containing prehensile-tailed Cyrtodactylus and have collapsed several clades in figure 6 to make the phylogeny easier to read. The base of the tree, including C. angularis, C. tibetanus, outgroups, and a clade of the Myanmar species, is not shown. Overall, our ND 2 phylogenies had high bootstrap support and posterior probabilities; it showed substantial congruence with the phylogeny of Wood et al. (2012). As for the COI gene, we found no evidence of a close relationship between Cyrtodactylus lateralis and C. elok using the ND 2 gene (Fig. 6). We found that C. elok and C. interdigitalis are closely related and belong to a clade containing the C. pulchellus Group, a large radiation of 18 Australo-Papuan species, and the mainland species C. oldhami, C. tygroides, C. hontreensis, C. intermedius, and C. peguensis. This result agrees with that of Wood et al. (2012), although they found the C. pulchellus Group to be sister to the C. elok + C. interdigitalis clade, whereas we found the mainland species to be sister to this pair. Nonetheless, C. elok and C. interdigitalis have relatively long branch lengths and an uncorrected genetic distance of 22.9 % from one another (Table 2). In contrast, Cyrtodactylus lateralis belongs to a large clade of mostly Sunda Shelf and Philippine species. Cyrtodactylus lateralis is sister to C. durio and their ND 2 sequences have diverged by 11.7 %. These two species have yet to show any clear affinitiess to three clades in the crown clade of figure 6, and we show all four lineages as an unresolved polytomy. The distant relationship of C. lateralis to the C. elok + C. interdigitalis clade is also reflected in high uncorrected genetic distances between C. lateralis and C. elok (27.6 %) and C. lateralis and C. interdigitalis (26.7 %)., Published as part of Harvey, Michael B., O'Connell, Kyle A., Wostl, Elijah, Riyanto, Awal, Kurniawan, Nia, Smith, Eric N. & Grismer, L. Lee, 2016, Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and Phylogeny of the Prehensile-tailed Cyrtodactylus, pp. 517-540 in Zootaxa 4107 (4) on pages 527-531, DOI: 10.11646/zootaxa.4107.4.3, http://zenodo.org/record/256578, {"references":["Uetz, P. & Hosek, J. (Eds.) (2015) The Reptile Database. Available from: http: // www. reptile-database. org (accessed 1 October 2015)","Grismer, L. L., Anuar, S., Quah, E., Muin, M. A., Onn, C. K., Grismer, J. L. & Ahmad, N. (2010) A new spiny, prehensile-tailed species of Cyrtodactylus (Squamata: Gekkonidae) from Peninsular Malaysia with a preliminary hypothesis of relationships based on morphology. Zootaxa, 2625, 40 - 52.","Ulber, T. (1993) Bemerkungen uber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen Arten (Reptilia: Gekkonidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin. Zoologisches Museum und Institut fur Spezielle Zoologie, 69, 187 - 200. http: // dx. doi. org / 10.1002 / mmnz. 19930690202","Linkem, C. W., McGuire, J. A., Hayden, C. J., Mohammed, I. S., Bickford, D. P. & Brown, R. M. (2008) A new species of bent-toe gecko (Gekkonidae: Cyrtodactylus) from Sulawesi Island, eastern Indonesia. Herpetologica, 64, 224 - 234. http: // dx. doi. org / 10.1655 / 07 - 040.1","Kraus, F. (2007) A new species of Cyrtodactylus (Squamata: Gekkonidae) from western Papua New Guinea. Zootaxa, 1425 (1), 63 - 68. http: // dx. doi. org / 10.11646 / zootaxa. 1425.1.8","Oliver, P., Edgar, P., Mumpuni, Iskandar, D. T. & Lilley, R. (2009) A new species of bent-toed gecko (Cyrtodactylus: Gekkonidae) from Seram Island, Indonesia. Zootaxa, 2115, 47 - 55.","Rosler, H. & Glaw, F. (2008) A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Malaysia including a literature survey of mensural and meristic data in the genus. Zootaxa, 1729, 8 - 22.","Wood, P. L. Jr., Heinicke, M. P., Jackman, T. R. & Bauer, A. M. (2012) Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution, 65, 992 - 1003. http: // dx. doi. org / 10.1016 / j. ympev. 2012.08.025"]}

Cyrtodactylus psarops sp. nov. Harvey, O���Connell, Barraza, Riyanto, Kurniawan & Smith Holotype. An adult female (MZB 9699; field number ENS 14612, Fig. 1) from Gunung Tanggamus, above Gisting, 5.424 �� S, 104.692 �� N, 1163 m, Kabupaten Tanggamus, Lampung Province, Sumatra, Indonesia, collected June 11, 2013, by M. B. Harvey, E. N. Smith, and field parties from BC, MZB, UB, and UTA. Paratypes (14). Four males (MZB 9686���9687, MZB 9700, UTA R- 61589) and one unsexed juvenile (UTA R- 61579) from forest above Ngarip, 5.280��� 5.284 �� S, 104.557��� 104.558 ��E, 1318���1419 m; four females (UTA R- 61580���61581, MZB 9690���9691) and one male (UTA R- 61590) from forest NE of Ngarip, 5.298- 5.314 �� S, 104.528 ���104.546, 950��� 1065 m; one male (MZB 9697) and two females (UTA R- 61582���61583) from Gunung Tanggamus, above Gisting, 5.422��� 5.424 �� S, 104.692��� 104.695 �� N, 1016���1163 m, Lampung, Sumatra, Indonesia; and one female (MZB 9700) from Gunung Pesagi (known locally as ���Masagi���), Remanan Jaya, Muara dua, Sumatera Selatan Province, Sumatra, Indonesia. M. B. Harvey, E. N. Smith, and field parties from BC, MZB, UB, and UTA collected the paratypes on 9���17 June 2013. Diagnosis. A medium-size species of Cyrtodactylus reaching 82 mm SVL and distinguished from all congeners by the following combination of characters: (1) body moderately robust; limbs and digits relatively long; (2) tail 1.06���1.35 times longer than body; (3) subpyramidal, weakly keeled tubercles extending from frontal region and supraorbital skin to base of tail, 23���26 irregular longitudinal rows of tubercles at midbody, 28���38 irregular transverse rows between limbs; (4) dorsal antebrachium and often brachium tuberculate; (5) tubercles on tail restricted to basal third; (6) 38���49 ventrals in a transverse row at midbody; (7) subcaudals not transversely enlarged; (8) deep, subtriangular depression in precloacal region of males; (9) 28���32 femoral and precloacal pores in males, arrayed in a continuous series (fewer pore primordia usually present in females), precloacal pores sunken into precloacal depression; (10) scale at anterior apex of pore-bearing scales greatly enlarged, sunken into precloacal depression in males, superficial in females; (11) pore-bearing series sharply transitioning to tiny granular scales of postaxial surface of thigh; (12) 18���22 lamellae under fourth toe; (13) cloacal tubercles usually single; (14) dorsal pattern of brown blotches edged first in dark brown to black then in pale gray or brown; at least some tubercles on flanks yellow to pale cream; (15) labials charcoal or dark brown with yellow or cream spots; occiput with few large blotches; long postocular stripe extending beyond arm onto flank; (16) 7���9 transverse, irregular or broken bands on body; 10���13 regular bands on tail, mostly complete ventrally. Description. Medium-size Cyrtodactylus reaching at least 82 mm SVL in females, 74 mm in males; smallest juvenile 42 mm SVL; body moderately robust, accounting for 64.9���74.3 % (n = 6) of SVL in males, 70.1���74.3 % (n = 8) of SVL in females; tail 1.06���1.35 (1.21 �� 0.10, n = 6) times longer than body and accounting for 51.6���57.4 % (54.7 �� 2.2, n = 6) of total length; head triangular in dorsal view; head 25.7���35.1 % (29.1 �� 3.2, n = 6) of SVL, 57.8���63.2 % (61.2 �� 2.2, n = 6) as wide as long, and 33.7 ���44.0% (37.7 �� 4.5, n = 6) as deep as wide in males and 27.3���29.8 % (28.7 �� 0.8, n = 8) of SVL, 54.5���62.4 % (58.4 �� 2.8, n = 8) as wide as long, and 31.7���40.6 % (36.6 �� 3.1, n = 8) as deep as wide in females. Rostral narrower than mental to as wide as mental (67.6 ���100.0%, 85.2 �� 8.7, n = 15, of width of mental), posteriorly forming border of nostril; dorsal margin of rostral deeply notched medially by shallow groove; rostral groove bifurcating slightly ventral to center of nostril; rostral bordered posteriorly by three (60 %, n = 15), four (27 %, n = 15), or five (13 %, n = 15) postrostrals; postrostral at midline projecting into posterior margin of rostral, except in UTA R- 61582 where posterior margin of rostral straight; internarial region narrow, 16.8���22.6 % (19.3 �� 1.8, n = 15) as wide as head; distance between anterior margins of orbit 30.3���38.6 % (35.3 �� 2.5, n = 15) as wide as head; shallow, key-hole shaped depression occupying most of prefrontal, frontal, and parietal areas (depression bordered laterally by slightly raised canthal, circumorbital, and supratemporal ridges and thickened skin at occiput); scales on dorsal snout granular, subpyrimidal, smooth, subequal in size, about as large as tubercles in parietal regions and distinctly smaller than granules between eyes; tubercles extending to frontal region and supraorbital skin; tubercles on dorsal surface of head subpyrimidal and smooth, much smaller than those on body, each surrounded by seven or eight granular scales; dorsal body tubercles unicarinate, oval, slightly projecting from dorsum, about five times as large as dorsal granules, 23���26 (23 �� 2, n = 14) longitudinal rows at midbody including scales in ventrolateral folds, 28���38 (33 �� 3, n = 14) irregular transverse rows between limbs; dorsal granules smooth. Nostril oval, directed postero-laterally and somewhat dorsally; bordered by rostral, postrostral, supranasal, nasal, and narrow antero-dorsal process of first supralabial; supranasal rounded to rhomboidal, about three times as large as adjacent postnasals; postnasals 5���7 (6 �� 1, n = 14); snout length 34.5���42.6 % (38.1 �� 2.1, n = 15) of head length and distance from orbit to nostril accounting for 73.4���80.3 % (77.0 �� 2.0, n = 15) of snout length; loreal region sloping; lips somewhat flared; 16���22 (20 �� 2, n = 15) loreals between nasal and anterior border of orbit; lorilabials in front of orbit flat, plate-like, enlarged relative to other loreals (1.5���3 times as large as loreals above them, but much smaller than supralabials); eye separated from supralabials by one or two longitudinal rows of tiny granules; diameter of orbit 19.0��� 25.7 % (23.3 �� 2.0, n = 15) of head length; temporal region with granular scales separating large, rounded, smooth or weakly keeled tubercles; distance from posterior border of orbit to anterior border of auditory meatus 24.1���33.5 % (29.3 �� 2.4, n = 15) of head length; auditory meatus deep, elliptical, directed postero-dorsally at about 40 �� to vertical, positioned at posterior end of head and with its ventral margin at same level as rictus; smallest diameter of auditory meatus 60���70 % smaller than greatest diameter; enlarged supralabials 9���12 (11 �� 1, n = 16), last separated by 1���5 small granular scales from rictus (12���16 total supralabials; 8���10 from center of orbit to rostral); first supralabial with antero-dorsal extension reaching nostril and separating lorilabials from rostral; remaining supralabials subrectangular with rounded dorsal edges, reducing in size to rictus. Mental triangular; infralabials 8���11 (10 �� 1, n = 16; 6���8 from center of orbit); pair of postmentals in contact medially; second pair of postmentals oval to squarish, about half as wide as first pair, separated from one another medially by 6���11 (9 �� 1, n = 15) small granular scales bordering first pair of postmentals; sublabial row decreasing in size posteriorly, not differentiated from adjacent gulars at level of eye; gulars small, smooth, granular, nonimbricate, much smaller than ventrals; in front of pectoral girdle, gulars grading into progressively larger, imbricate scales; scale of chest like ventrals; ventrals smooth, imbricate, flat, circular, about as wide as tubercles on dorsum; ventrals in transverse row at midbody 38���49 (44 �� 4, n = 13); ventrolateral skin fold distinct, containing tubercles. Tail subcylindrical, though somewhat flattened dorsally and ventrally; tubercles restricted to 16.0��� 24.9 % (19.1 �� 3.1, n = 6) of intact tails; distribution of tubercles at base of tail like dorsal body, distally tubercles not projecting and arrayed in 3���6 verticils; verticils separated by 6���9 small scales along dorsal midline; tubercles on tail not extending to ventral side, although some scales of verticils may be slightly enlarged relative to other subcaudals; subcaudals flat, homogenous, 1.5���2 times as large as scales on sides of tail; each subcaudal about as long as wide; transversely enlarged subcaudal plates absent; cloacal tubercles rounded, smooth, single, slightly smaller than dorsal tubercles; tip of tail capped by granular scales. Brachium covered in small, homogenous, weakly keeled scales; brachial tubercles usually present (67 %, n = 15); antebrachium tuberculate; antebrachial tubercles subpyrimidal, unicarinate, about four times as large as granular scales between them, slightly smaller than those on dorsum; ventral surfaces of brachium and antebrachium covered in smooth, slightly imbricate, homogenous scales; scales on palm and dorsal surface of hand smooth, imbricate; scales under finger I 8���11 (n = 15 for all digits), under finger II 13���15, under finger III 14���18, under finger IV 16���19, under finger V 13���16 (all counts excluding ungual scale); one or two scales widened and similar to lamellae extending onto palm at base of digits. Thigh, shank, and postaxial half of dorsal foot (at base of toes IV���V) tuberculate; tubercles of leg subpyrimidal, unicarinate, about same size as those on dorsum, about five times as large as small granular scales between them; preaxial and ventral surface of thigh covered in rounded, flat, imbricate scales; ventral thigh with enlarged row of pore-bearing scales and similarly enlarged row of scales anterior to pore-bearing row; this doubled row of scales arranged in an inverted V with its apex at midline of belly; abrupt transition to tiny granules on postaxial surface of thigh; postfemoral granules only one-fifth to one-sixth size of pore-bearing scales; sole covered in flat, imbricate, rounded scales; scales under toe I 8���11 (n = 15), under toe II 13���15, under toe III 16���20; under toe IV 18���22, under toe V 17���21; one or two scales widened and similar to lamellae extending onto sole at base of digits. Arrangement of pore-bearing scales sexually dimorphic; both sexes with single, greatly enlarged scale at apex of pore-bearing series (Fig. 2); apical scale twice as long as wide, with single pore or pore primordium at its posterior end; apical scale superficial in females and juveniles, sunken into precloacal depression of males; total pores 28���32 (30 �� 2, n = 6) in males; pore rows of males continuous, extending onto thighs; pore primordia present (75 %, n = 8) or absent (25 %) in females; when present, total pore primordia 13���29 (20 �� 6, n = 6); in two specimens distal primordia separated by gaps of 2 / 4 and 3 / 3 (right/left) scales on thigh; in specimens without gaps, primordia extending for short distance onto thigh (50 %, n = 6) or restricted to body (50 %). Coloration. In life (based on field notes of M.B. Harvey and photos of all specimens in the type series), Cyrtodactylus psarops has a gray, greenish gray, or greenish brown ground coloration with brown markings widely edged in black. On the flanks, tubercles edging the brown dorsal markings are frequently bright yellow, sharply contrasting with the adjacent color. Anastomosing brown and pale yellow markings cover the dorsal snout. The supraciliaries are also yellow with some black areas. The rostral, mental, narial region, and labials are mostly charcoal to black, and the labials are marked by widely spaced, distinctive yellow spots. Dorsally, several large subcircular blotches cover the parietal area. A wide postocular stripe extends beyond the arm onto the flank. A transverse nuchal band may partially fuse to the postocular stripe, although the two markings are usually separate from one another. The iris is reddish brown, edged in black and with a black reticulum. The ciliary body is only slightly lighter than the iris and indistict. In Cyrtodactylus psarops, the distal third of the tongue is black above and below, whereas the rest of the tongue is pink. Cyrtodactylus psarops (Fig. 3) has one band on the neck, 7���9 (8 �� 1, n = 15) bands on the body, and 10���13 (11 �� 1, n = 6) bands on the tail. Bands on the neck and body frequently break up into subcircular blotches. Bands on the tail are narrower than the pale interspaces at the base of the tail but longer than the interspaces distally. Distally, the caudal interspaces become increasingly more pale. Ventrally the caudal bands are continuous (i.e., the tail is ringed). Regenerated tails are entirely tan above and below. The brachium, antebrachium, thigh, and shank each bears three to four bands. Banding extends onto the hand and foot, although the digits are mostly charcoal or dark brown. Except for the tail, the venter is pale pink to purple. Palms and soles are heavily melanic, whereas the porebearing scales are distinctly paler than scales around them. Adult males have a distinctive chalky white, semilunar area anterior and lateral to the precloacal depression. This chalky area extends to the groin. Etymology. This species��� name psarops is a noun in apposition derived from the Greek words psaros, meaning speckled, and ops, meaning face. The new name alludes to the characteristic yellow speckling of the labials and snout in this species. Standard English name. Speckle-faced Bent-toed Geckos. Standard Indonesian name. Cicak Jari Lengkung Muka Bintik. Distribution and natural history. We collected most of the type specimens from humid forest, coffee plantations, and adjacent secondary growth on Gunung Tanggamus and mountains surrounding Ngarip, a small town in the southern Bukit Barisan Range, Lampung, Sumatra (Fig. 4). Cyrtodactylus psarops extends into adjacent Sumatera Selatan at Gunung Pesagi. We encountered most specimens on the forest floor at night (19:00��� 21: 10 hrs), however this species readily ascends into low vegetation. We found UTA R- 61581 about 2 m aboveground on the stem of a plant at the forest edge and MZB 9702 about 1 m aboveground on a dead tree in a coffee plantation. Cyrtodactylus psarops occurs between 950���1419 m, however, its elevational range may be broader. We did not visit sites at higher or lower elevations in the vicinity of the type locality., Published as part of Harvey, Michael B., O'Connell, Kyle A., Barraza, Gabriel, Riyanto, Awal, Kurniawan, Nia & Smith, Eric N., 2015, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the Southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny, pp. 495-516 in Zootaxa 4020 (3) on pages 498-503, DOI: 10.11646/zootaxa.4020.3.5, http://zenodo.org/record/240877

Cyrtodactylus semicinctus sp. nov. Harvey, O���Connell, Barraza, Riyanto, Kurniawan & Smith Holotype. An adult male (MZB 9703, collector���s tag ENS 14958, Fig. 5) from montane forest on a trail above Kayu Aro, leading to the top of Mt. Kerinci, Kabupaten Kerinci, Jambi Province, Sumatra, Indonesia, 1905 m, 1.74159 �� S, 101.26015 ��E, collected on 24 June 2013 by M. B. Harvey and K. A. O���Connell. Paratypes (7). Two males (MZB 9695, 9696), four females (MZB 9694, UTA R- 61576, 61577, 61588), and one unsexed juvenile (MZB 9701) collected from the same trail as the holotype, 1752���2006 m, on 21 June 2014 by M. B. Harvey, E. N. Smith, and other members of the summer 2013 field party. Diagnosis. A medium size species of Cyrtodactylus reaching 89 mm SVL and distinguished from all congeners by the following combination of characters: (1) body moderately robust; limbs and digits relatively long; (2) tail 1.05���1.19 times longer than body; (3) subpyramidal, weakly keeled tubercles extending from parietal region and supraorbital skin to base of tail, 24���27 irregular longitudinal rows of tubercles at midbody, 29���35 irregular transverse rows between limbs; (4) dorsal antebrachium tuberculate, brachium lacking tubercles; (5) tubercles on tail restricted to basal third; (6) 33���44 ventrals in a transverse row at midbody; (7) subcaudals not transversely enlarged; (8) deep, subtriangular precloacal depression in males; (9) 36���38 femoral and precloacal pores in males, arrayed in a continuous series (fewer pore primordia usually present in females), sunken into precloacal depression; (10) scale at anterior apex of pore-bearing scales greatly enlarged, sunken into precloacal depression in males, superficial in females; (11) pore-bearing series sharply transitioning to tiny granular scales of postaxial surface of thigh; (12) 19���22 lamellae under fourth toe; (13) cloacal tubercles 2 / 2 or 3 / 3 contacting one another; (14) dorsal pattern of brown blotches widely edged in black; tubercles on flanks yellow to pale cream; (15) labials mostly yellow; wide U-shaped band around occiput to loreal region; (16) 5���7 transverse, irregular or broken bands on body; 9���11 regular bands on tail, incomplete ventrally, except near distal tip of some specimens. Description. Medium-size Cyrtodactylus reaching at least 89 mm SVL in females, 75 mm in males; smallest juvenile 36 mm SVL; body moderately robust, accounting for 69.9���72.9 % (n = 3) of SVL in males, 70.1���74.3 % (n = 4) of SVL in females; tail 1.05���1.19 (1.10 �� 0.05, n = 5) times longer than body and accounting for 51.4���54.4 % (52.4 �� 1.2, n = 5) of total length; head triangular in dorsal view; head 27.1���30.1 % of SVL, 62.4���71.1 % as wide as long, and 37.6���40.3 % as deep as wide in males (n = 3) and 25.7���29.9 % of SVL, 58.7���72.3 % as wide as long, and 32.7���37.1 % as deep as wide in females (n = 4). Rostral narrower than mental (76.7���99.5 %, 89.5 �� 8.6, n = 8, of width of mental), posteriorly forming border of nostril; dorsal margin of rostral deeply notched medially by shallow groove; rostral bordered posteriorly by three (50 %, n = 8), four (38 %, n = 8), or five (12 %, n = 8) postrostrals; postrostral at midline projecting into posterior margin of rostral; internarial region narrow, 17.2���21.1 % (19.5 �� 1.4, n = 8) as wide as head; distance between anterior margins of orbit 29.2���39.6 % (34.8 �� 3.9, n = 8) as wide as head; shallow, key-hole shaped depression occupying most of prefrontal, frontal, and parietal areas (depression bordered laterally by slightly raised canthal, circumorbital, and supratemporal ridges and thickened skin at occiput); scales on dorsal snout granular, subpyrimidal, smooth, subequal in size, and slightly larger than granules of frontal and parietal regions; tubercles extending to parietal region and on supraorbital skin but absent from snout and depressed areas between eyes; tubercles on dorsal surface of head subpyrimidal and smooth, much smaller than those on body, each surrounded by seven or eight granular scales; dorsal body tubercles unicarinate, oval, slightly projecting from dorsum, about five times as large as dorsal granules, 24���27 (25 �� 1, n = 8) longitudinal rows at midbody including scales in ventrolateral folds, 29���35 (32 �� 2, n = 8) irregular transverse rows between limbs; dorsal granules smooth. Nostril oval, directed postero-laterally and somewhat dorsally; bordered by rostral, postrostral, supranasal, nasal, and narrow antero-dorsal process of first supralabial; supranasal rounded, about twice as large as adjacent postnasals; postnasals 4���6 (5 �� 0, n = 16); snout length 37.3���44.7 % (40.8 �� 2.1, n = 8) of head length and distance from orbit to nostril accounting for 68.1���77.6 % (72.7 �� 2.8, n = 8) of snout length; loreal region sloping; lips flared; 16���23 (18 �� 2, n = 8) loreals between nasal and anterior border of orbit; lorilabials in front of orbit flat, plate-like, enlarged relative to other loreals (1.5���2 times as large as loreals above them, but much smaller than supralabials); eye separated from supralabials by one or two longitudinal rows of tiny granules; diameter of orbit 20.8���26.6 % (23.4 �� 2.0, n = 8) of head length; temporal region with granular scales separating large, subpyrimidal, and keeled tubercles; distance from posterior border of orbit to anterior border of auditory meatus 27.0��� 33.3 % (29.7 �� 2.3, n = 8) of head length; auditory meatus deep, narrowly elliptical, directed postero-dorsally at about 30 �� to vertical, positioned at posterior end of head and with its ventral margin at same level as rictus; smallest diameter of auditory meatus 60���70 % smaller than greatest diameter; enlarged supralabials 8���11 (10 �� 1, n = 15), last separated by 2���4 small granular scales from rictus (10���15 total supralabials; 7���9 from center of orbit to rostral); first supralabial with antero-dorsal extension reaching nostril and separating lorilabials from rostral; remaining supralabials subrectangular with rounded dorsal edges, reducing in size to rictus. Mental triangular; infralabials 8���10 (9 �� 1, n = 15; 6���7 from center of orbit); pair of postmentals in contact medially; second pair of postmentals squarish, about half as wide as first pair, separated from one another medially by 6���8 (7 �� 1, n = 8) small granular scales bordering first pair of postmentals; sublabial row decreasing in size posteriorly; gulars small, smooth, granular, nonimbricate, much smaller than ventrals; gulars grading into progressively larger, imbricate scales just in front of pectoral girdle; scale of chest like ventrals; ventrals smooth, imbricate, flat, circular, slightly smaller than tubercles on dorsum, but much smaller than dorsal granules; ventrals in transverse row at midbody 33���44 (38 �� 5, n = 6); ventrolateral skin fold distinct, containing tubercles. Tail subcylindrical, though somewhat flattened dorsally and ventrally; tubercles restricted to 17.2���23.3 % of intact tails (n = 2); distribution of tubercles at base of tail like dorsal body, distally tubercles not projecting and arrayed in 1���5 verticils; verticiles separated by 5���7 small scales along dorsal midline; tubercles on tail not extending to ventral side, although some scales of verticils may be slightly enlarged relative to other subcaudals; subcaudals flat, homogenous, two or three times as large as scales on sides of tail; each subcaudal about as long as wide; transversely enlarged subcaudal plates absent; in specimens with regenerated tails, subcaudals just proximal to regenerated portion enlarged: about half again as long as scale row proximal to them; cloacal tubercles rounded, smooth, two (88 %, n = 16) or three (12 %, n = 16, both counts of three in one specimen: UTA R- 61577), in contact with one another, half again as large as dorsal tubercles; tip of tail capped by granular scales. Brachium covered in small, homogenous, weakly keeled scales; brachial tubercles absent; antebrachium tuberculate; antebrachial tubercles subpyrimidal, unicarinate, about four times as large as granular scales between them, slightly smaller than those on dorsum; ventral surfaces of brachium and antebrachium covered in smooth, slightly imbricate, homogenous scales; scales on palm and dorsal surface of hand granular, smooth, weakly (ventrally) to moderately (dorsal surface) imbricate; scales under finger I 8���10 (n = 8 for all digits), under finger II 12���15, under finger III 15���17, under finger IV 16���18, under finger V 12���15 (all counts excluding ungual scale); one or two scales widened and similar to lamellae extending onto palm at base of digits. Thigh, shank, and postaxial half of dorsal foot (at base of toes IV���V) tuberculate; tubercles of leg subpyrimidal, unicarinate, about same size as those on dorsum, about five times as large as small granular scales between them; preaxial and ventral surface of thigh covered in rounded, flat, imbricate scales; ventral thigh with enlarged row of pore-bearing scales and similarly enlarged row of scales anterior to pore-bearing row; this doubled row of scales arranged in an inverted V with its apex at midline of belly; abrupt transition to tiny granules on postaxial surface of thigh; postfemoral granules only one-fifth to one-sixth size of pore-bearing scales; sole covered in flat, imbricate, rounded scales; scales under toe I 8���11, under toe II 12���15, under toe III 17���19; under toe IV 19��� 22, under toe V 17���20; one or two scales widened and similar to lamellae extending onto sole at base of digits. Arrangement of pore-bearing scales sexually dimorphic; both sexes with single, greatly enlarged scale at apex of pore-bearing series; apical scale twice as long as wide, with single pore or pore primordium at its posterior end; apical scale superficial in females and juveniles, sunken into precloacal depression in males; total pores 36 and 38 in two adult males (SVL 75 mm and 68 mm, respectively), only 14 in small male (SVL 55 mm) perhaps due to ontogenetic differences; two females (SVL 51 mm and 89 mm, respectively) with 17 and 19 pore primordia, one lacking primordia (SVL 83 mm); in both sexes pores and/or primordia extending onto thigh. Coloration. In life, Cyrtodactylus semicinctus has a light gray, greenish tan, or dull orange ground coloration with brown to reddish brown markings widely edged in black (MZB 9695 was in the process of shedding and its ground coloration was gray rather than orange or greenish tan). Narrow markings are entirely black. The dorsal tubercles are a paler shade than the dorsal ground color. On the flanks, even some of the tubercles within dark markings are this pale color, so that the flanks appear to be densely speckled. A yellow wash covers the labials, rostral, and supraciliary scales. These yellowish scales are mostly immaculate, although small, irregular black blotches mark the narial and internarial region and sometimes extend ventrally onto the rostral and or first supralabial. Dorsally, the head usually has one to three poorly defined blotches in the parietal region and a sharp black line edging the gray to yellow supraciliaries medially. This species has a distinctive U-shaped marking extending from the loreal regions around the occiput. The marking begins a short distance behind the nasal just above the enlarged series of lorilabials and widens just in front of the eye. Behind the eye and around the occiput, a slightly more narrow pale band edges the U-shaped band dorsally and accentuates it. The iris is reddish brown to greenish bronze, edged in black and with a black reticulum. The ciliary body is only slightly lighter than the iris and indistict. Cyrtodactylus semicinctus (Fig. 6) has one band on the neck, 5���7 (6 �� 1, n = 8) bands on the body, and 9���11 (10 �� 1, n = 5) bands on the tail. Bands on the neck and body are jagged, irregular, and frequently broken on the flanks. When broken, blotches on the flanks may fuse with one another, sometimes to produce long sinuous markings. In particular, the band on the neck usually breaks up into blotches. However, the band on the neck does not fuse to the U-shaped nuchal band. Bands on the tail are slightly longer than the pale interspaces. Except near the distal tip of the tail of some specimens, the bands are not continuous ventrally (i.e., the tail is banded, not ringed; all specimens have at least the basal one-third of the tail not regenerated and none have ringed tails). Distally, the caudal interspaces become increasingly more pale. Regenerated tails are entirely black or mostly black with few scattered cream scales. The brachium, antebrachium, thigh, and shank each bears two or three jagged, irregular bands. Except for the tail, the venter is mostly rose pink to grayish purple. Palms and soles are a darker shade of the same ground color, whereas the pore-bearing series are distinctly paler than scales around them. Some scales at the edges of the venter are the same color as the dorsal ground coloration. They form a diffuse reticulum against the rose pink to grayish purple ground color. This diffuse, greenish tan to orange reticulum is restricted to the lateral edges of the belly and pre- and postaxial margins of the ventral limbs, but extends across the entire gular region. The cloacal tubercles are greenish tan or orange. Ventrally, intact tails are tessellated with charcoal and elements of the dorsal ground coloration. Etymology. This species��� name semicinctus is a masculine adjective derived from the Latin noun cinctum, meaning girdle or belt, and prefix semi, meaning partial or incomplete. The name alludes to the incomplete caudal rings of this species. Standard English name. Kerinci Bent-toed Geckos. Standard Indonesian name. Cicak Jari Lengkung Kerinci. Distribution and natural history. This species is known only from the type locality. We collected the entire type series between 1752���2006 m along a trail above the tiny hamlet of Kayu Aro. The trail extends through apparently pristine forest to the top of Mt. Kerinci and is frequented by hikers. Field parties collected Cyrtodactylus semicinctus on the forest floor at night (1950���2340 hrs) and we found most specimens around tree trunks. In field notes, Harvey noted that UTA R- 61576 emitted a faint ���baaap��� when first grasped and a louder ���raaank��� when struggling as the senior author collected color notes., Published as part of Harvey, Michael B., O'Connell, Kyle A., Barraza, Gabriel, Riyanto, Awal, Kurniawan, Nia & Smith, Eric N., 2015, Two new species of Cyrtodactylus (Squamata: Gekkonidae) from the Southern Bukit Barisan Range of Sumatra and an estimation of their phylogeny, pp. 495-516 in Zootaxa 4020 (3) on pages 503-507, DOI: 10.11646/zootaxa.4020.3.5, http://zenodo.org/record/240877