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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."]}

This file contains supplementary text and 14 supplementary figures. The supplementary text provides (i) supplementary morphological descriptions; (ii) methodology on estimating squamate body-sizes; (iii) interpretations of the palaeoenvironmental setting of Magnuviator ovimonsensis and other relevant fossil iguanomorphs; (iv) new systematic interpretations of Late Cretaceous lizards previously assigned to Pleurodonta; (v) cladistic analysis methodologies and results; (vi) character scorings of Magnuviator ovimonsensis and Pariguana lancensis; (vii) information regarding CT and µCT scans; and (viii) a complete strict consensus tree and apomorphy list derived from our morphology-based cladistic analysis. The supplementary figures show additional specimen images, cladograms, and comparative body-size estimates of fossil and extant lizards. The text provides background information for interpretations made in the manuscript.

Head, Jason J., Barrett Fls, Paul M., Rayfield, Emily J. (2009): Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae). Zoological Journal of the Linnean Society 155 (2): 445-457, DOI: 10.1111/j.1096-3642.2008.00448.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00448.x