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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."]}

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 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 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 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 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."]}

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

Catastrophic events, such as volcanic eruptions, can have profound impacts on the demographic histories of resident taxa. Due to its presumed effect on biodiversity, the Pleistocene eruption of super-volcano Toba has received abundant attention. We test the effects of the Toba eruption on the diversification, genetic diversity, and demography of three co-distributed species of parachuting frogs (Genus Rhacophorus ) on Sumatra. We generate target-capture data (∼950 loci and ∼440,000 bp) for three species of parachuting frogs and use these data paired with previously generated double digest restriction-site associated DNA (ddRADseq) data to estimate population structure and genetic diversity, to test for population size changes using demographic modeling, and to estimate the temporal clustering of size change events using a full-likelihood Bayesian method. We find that populations around Toba exhibit reduced genetic diversity compared with southern populations, and that these northern populations exhibit a signal of contraction around the time of the eruption (∼80 kya). However, we infer a stronger signal of expansion in southern populations around ∼400 kya, and at least two of the northern populations may have also expanded at this time. Taken together, this suggests that the Toba eruption precipitated population declines in northern populations, but that the demographic history of these three species was more greatly impacted by mid-Pleistocene forest expansion, supporting local rather than regional effects of the Toba eruption.

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

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