Craugastor, Cope, 1862

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, Herpetological Monographs 36 (1), pp. 1-48 : 38-40

DICHOTOMOUS KEY TO THE CRAUGASTOR MEXICANUS SERIES

1a. Large or medium-sized adults (females> 20 mm SVL, males> 16 mm SVL); axillary gland present or absent; supratympanic fold usually developed (except for moderately to poorly developed in C. montanus ); both vomerine odontophores and posterolateral projection of frontoparietal present (except for some C. montanus with no odontophores [but with frontoparietal projection], and some C. mexicanus with no frontoparietal projection [but with vomerine odontophores]) ........... 2

1b. Small sized adults (females <20 mm SVL, males <16 mm SVL); axillary gland always absent; supratympanic fold absent or poorly developed; with no vomerine odontophores (except in C. candelariensis ) ................ 5

2a. Heelnotreachingnostrilwhen legadpressedtobody; two or three metacarpal tubercles (including thenar; outer smallest, middle largest); outer/inner metatarsaltubercle ratio 0.40–0.55; testis usually unpigmented (sometimes lightly pigmented); ovaries unpigmented; fromthe highlands of Guerrero and central Oaxaca (1900–2600 m elevation) ............. Craugastor omiltemanus (Günther)

2b. Heel reaching tip of snout or beyond when leg adpressed to body; always three metacarpal tubercles (including thenar); outer/inner metatarsaltubercle ratio variable (0.36–0.75); testis strongly pigmented; ovaries always pigmented (strongly or lightly) ............ 3

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3a. Medium-sized adult females (<16 mm SVL); crus 53– 59% SVL; with or without vomerine odontophores; supratympanic fold moderately to poorly developed; labium always barred; middle metacarpal tubercle largest and outer very small; outer tarsal heel tubercles Ĺ 2 (close to heel); ovaries pigmented; highlands east of the Isthmus of Tehuantepec (from the Sierra Madre del Sur of eastern Chiapas and western Guatemala) ................................... Craugastor montanus (Taylor)

3b. Large-sized adult females (> 16 mm SVL); crus 53– 73% SVL; always with vomerine odontophores; supratympanic fold well-developed; labium variable in pattern; middle metacarpal tubercle about same size as thenar, outer smaller; outer tarsal heel tubercles 0– 5; ovaries pigmented or not; highlands west of the Isthmus of Tehuantepec (from Oaxaca and Guerrero to Puebla and Hidalgo, including the central highlands of Veracruz) ......................................................... 4

4a. Crus>62% SVL (62–73%); heel always beyond tip of snout when leg adpressed to body; outer/inner metatarsal tubercle ratio (0.36[0]–0.58); ovaries unpigmented or only slightly pigmented; from the Sierra Madre del Sur of Guerrero west of Chilpancingo (1900–2850 m elevation) .................... Craugastor saltator (Taylor)

4b. Crus <69% SVL (53–69%); heel at snout tip or sometimes beyond when leg adpressed to body; outer/inner metatarsaltubercle ratio (0.50–0.75); ovaries pigmented or not; from Guerrero east of Chilpancingo to Oaxaca and to the north in Puebla, Hidalgo and the adjacent Veracruz highlands (1500– 3015 m elevation) ...... Craugastormexicanus (Brocchi)

5a. Vomerine odontophores present; outer two fingers and toes lanceolate to mucronate; body flank unicolor pale; no raised eyelid tubercles; gular region not pigmented or only lightly and uniformly peppered; face never barred; postrictal tubercles 2; metacarpal tubercles about equal in size; outer tarsal ridge with 3–8 tiny and pointed tubercles on slightly raised fringe; posterolateral projection of frontoparietal absent; from the Sierra Madre del Sur of Oaxaca, mountains above Puerto Escondido and Puerto Angel (668–1350 m elevation)............... Craugastor candelariensis Jameson et al.

5b. Vomerine odontophores absent; fingers and toes usually round or at most, slightly lanceolate; body flank darker anteriorly (toward axilla, except uniformly pale in some C. cueyatl ); usually with raised eyelid tubercles; gular region blotched or heavily peppered (rarely lightly peppered in C. pygmaeus ); face barred or not; postrictal tubercles 1 or 2; outer metacarpal tubercle usually longer and/or wider than inner (except about equal in size in C. cueyatl and C. pygmaeus ); outer tarsal ridge with no raised fringe and variable number of tubercles (0–7), flat and round (sometimes slightly pointed); posterolateral projection of frontoparietal present or absent ................................................. 6

6a. Face barred, one or two prominent bars below eye; no distinctive canthal stripe; one distinctive dark and rectangular blotch under lower jaw and at level of canthus (prominent in ventral view); gular region with pale spotting; finger tips not expanded; outer tarsal ridge with 0–4 extremely small, flat and round tubercles; posterolateral projection of frontoparietal present; from the Sierra Madre del Sur of Oaxaca in the mountains above Puerto Escondido (1245–1585 m elevation) ............ Craugastor polaclavus Jameson et al.

6b. Face barred or not, bars below eye variable; canthal stripe present or absent; no distinctive rectangular blotches under lower jaw and at level of canthus (not prominent in ventral view); gular region variable but, usually spotted or peppered; finger tips expanded or not; outer tarsal ridge with 0–7 extremely small, flat, and round to slightly pointed tubercles; posterolateral projection of frontoparietal present or absent; ........... 7

7a. Finger and toe tips round and conspicuously expanded; face barred, never with a canthal stripe; postrictal tubercles 1 (sometimes look like two fused); dorsal pattern two-toned or striped, and with a pale middorsal stripe; metacarpal tubercles about equal in size; outer tarsal ridge smooth and lacking tubercles; posterolateral projection of frontoparietal absent; from the lowlands of southern Mexico, from Puebla and Veracruz on the Gulf versant and southern Guerrero on the Pacific versant to Escuintla, Chiapas and possibly adjacent Guatemala (0–1580 m elevation)................................ Craugastor pygmaeus (Taylor)

7b. Finger tips round or lanceolate, not expanded or only slightly; toe tips slightly expanded to expanded, sometimes slightly lanceolated; face barred or with a pale supralabial stripe, canthal stripe present or absent (some C. bitonium and C. hobartsmithi ); postrictal tubercles 1 or 2; dorsal pattern variable; outer tarsal ridge smooth or tuberculated ( C. portilloensis and C. rubinus ) or always tuberculated; posterolateral projection of frontoparietal present or absent....................... 8

8a. Always with a canthal stripe and softly barred lips; dorsum uniformly dark; postrictal tubercles always 1, on each side; body flank uniformly pale or slightly darker anteriorly; gular region with evenly distributed fine melanocytes (evenly peppered); metacarpal tubercles about equal in size; finger tips round and not expanded; outer tarsal ridge with 3–7 rounded tubercles; posterolateral projection of frontoparietal present; from the slopes facing the Balsas River depression in the states of Mexico and Morelos (1670–2311 m elevation) ................. Craugastor cueyatl Jameson et al.

8b. Canthal stripe present or absent, and with soft or prominent bars on lips or with a supralabial pale stripe; dorsum uniformly pale, two-toned, or striped, never uniformly colored and darker than flanks; postrictal tubercles 1 or 2; body flank always darker anteriorly; gular region variable but usually melanocyte distribution forming light spotting (particularly under lower jaws and mid-gular area); outer metacarpal tubercle slightly larger than inner; finger tips not or just barely expanded, round or slightly lanceolated; outer tarsal ridge with 0–7 tubercles; posterolateral projection of frontoparietal present or absent ................................... 9

9a. Canthal stripe present or absent, and with bars on lips or a supralabial pale stripe; dorsum usually two-toned or striped; gular region with small melanocyte distribution forming a few small light spots under lower jaws and mid-gular area; finger tips slightly expanded, round or slightly lanceolated ( C. bitonium ); outer tarsal ridge with 1–6 tubercles, round to slightly pointed; posterolateral projection of frontoparietal absent ................. 10

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9b. Canthal stripe always present, and with bar(s) on lips; dorsum uniformly pale (as compared with flanks, one specimen of C. rubinus with traces of ill-defined and light blotching); gular region with melanocyte distribution forming prominent light spots under lower jaws and mid-gular area; finger tips usually not expanded (somewhat expanded in one specimen of C. rubinus ), round, never lanceolated; outer tarsal ridge with 0–4 tubercles, extremely small, flat, and round; posterolateral projection of frontoparietal present.................... 11

10a. Postrictal tubercles 1, on each side; finger and toe tips slightly lanceolated; finger lengths III> IV ± II> I; toe lengths always IV> III> V> II> I; outer/inner metatarsal tubercle ratio 0.40–0.75; not less than four outer tarsal tubercles, 4–6; from the Sierra Madre del Sur de Guerrero, south of Yextla (2071 m elevation)....................... Craugastor bitonium Jameson et al.

10b. Postrictal tubercles 1–2, on each side; finger and toe tips rounded, not lanceolated; finger lengths always III> IV> II> I; toelengths IV> III ± V> II> I; outer/inner metatarsal tubercle ratio 0.62–0.67; outer tarsal tubercles 1–6; from near Uruapan, Michoacán (1450–1700 m elevation) ................... Craugastor hobartsmithi (Taylor)

11a. Eyelid raised tubercles present or absent; postrictal tubercles 1–2, on each side; toe tips round and slightly expanded; finger lengths III> IV> II> I; toe lengths IV> III ± V> II> I; outer/inner metatarsaltubercle ratio 0.67–1.25; from the Sierra Madre del Sur of Oaxaca at Portillo del Rayo (1550–1585 m elevation).................. Craugastor portilloensis Jameson et al.

11b. Eyelid raised tubercles present; postrictal tubercles 1, on each side; toe tips slightly lanceolated and barely expanded; finger lengths III> IV ¼ II> I; toe lengths always IV> III> V> II> I; outer/innermetatarsal tubercle ratio 0.50–0.55; from the Pacific slopes of Jalisco west of Talpa de Allende (1771 m elevation)......................... Craugastor rubinus Jameson et al.