Neotropical Hylid Frogs, Genus Smilisca Part 6

=Guanacaste=: Tilaran, KU 36805-7; 8 km. NE Tilaran, KU 36803-4.

=Heredia=: Barranca del Rio Sarapiqui below Isla Bonita, KU 64282-3; Cariblanco, KU 32256-60, 41094 (skeleton), 64284, MCZ 7967; Isla Bonita, KU 32250-4; 4.2 km. W Puerto Viejo, KU 64285, 68485; 7.5 km. W Puerto Viejo, KU 68486; 1 km. S Puerto Viejo, KU 86518.

=Limon=: Bambu, USC 7182 (4); Batan, UMMZ 118582; Coen, MCZ 9825; La Lola, KU 32262-4, UF 4029, UMMZ 117678 (3); Los Diamantes, CNHM 101295-8, KU 25723-4, 32265, 64267-9; Pandora, UMMZ 122650 (2), USC 7188 (3), 7190; Puerto Limon, KU 32261; Rio Lari at Rio Dipari, 21 km. SW Amubre, USC 7177; Rio Toro Amarillo, 7 km. W Guapiles, KU 86519, 87683 (tadpoles); Suretka, KU 36808-10, 37168.

=Puntarenas=: Agua Buena, KU 36790; 1.6 km. E Buenos Aires, UMMZ 117578; 3 km. NW Buenos Aires, KU 64304; 4 km. N, 15 km. W Dominical, KU 68491-2 (tadpoles); Esparta, MCZ 8029-30, 8032; Golfito, KU 32270; 6 km. E Golfito, KU 84999-500 (skeletons); Gromaco, UMMZ 123677 (4); Palmar, KU 32269; 4 km. ESE Palmar Sur, KU 64305-6; 5.6 km. SE Palmar Sur, KU 68489 (tadpoles); 7.0 km. SE Palmar Sur, KU 68490 (young); 8.5 km. SE Piedras Blancas, KU 64292-303; Quebrada Boruca, 22 km. E Palmar Norte, KU 64291; Rincon, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), USC 7254; Rio Ferruviosa, 7 km. S Rincon, USC 7235; 1.6 km. WNW Villa Neily, KU 68493 (young), 68494 (tadpoles).

=San Jose=: San Isidro el General, KU 32249, UMMZ 75025; 10 km. N San Isidro el General, MCZ 29099-103; 13 km. WSW San Isidro el General, KU 86517; 15 km. WSW San Isidro el General, KU 68487 (tadpoles), 68488 (young), 68495 (young); 20 km. WSW San Isidro el General, KU 32248.

PANAMA: No province: Cano Saddle, USNM 69588; Punta de Pena, USNM 38733; Quipo, AMNH 18925-6. =Bocas del Toro=: Almirante, KU 80080, 91835-6; 1.6 km. W Almirante, KU 91837; 3 km. W Almirante, KU 91824 (skeleton), 91838-43, 91906-7; 11 km. NW Almirante, CNHM 67853-61; 13 km. W Almirante, KU 91825-7 (skeletons), 91844-9; Fish Creek, KU 92329; Isla Popa, KU 91850-1. =Ca.n.a.l Zone=: Barro Colorado Island, CNHM 6007, 13316, 13325, 13331, 13360-2, 13377-8, MCZ 24191-5, UF 7523, UMMZ 63547-60, 64457, 69497 (3); 3.7 km. W Cocoli, KU 67916; Fort Sherman, MCZ 10139; Gatun, MCZ 35644; Junction roads C25B and C16, TNHC 23839; Madden Forest Preserve, TNHC 23837-8. =Cocle=: El Valle, KU 77521-4, 77649 (tadpole), TNHC 23369. =Comarca del Baru=: Progreso, UMMZ 61085-9.

Colon: Achiote, KU 77516-20, 77648 (young); Rio Candelaria, CNHM 67851-2. =Darien=: Rio Esnape, Sambu Valley, MCZ 8539; Rio Sucubti, Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, Camp Townsend, AMNH 40988. =Panama=: NW slope Cerro Prominente, KU 80459; Finca La Sumbadora, KU 91914 (skeleton). =Chiriqui=: 2 km. W Concepcion, AMNH 68910.

COLUMBIA: =Antioquia=: Puerto Berrio, CNHM 30805 (Goin); Turbo, USNM 39899. =Caldas=: Pueblorrica, Santa Cecilia, CNHM 54768-71 (Goin).

=Choco=: No specific locality, AMNH 3984-6; Andagoya, BMNH 1915.10.21.

69-70, CNHM 81857 (Goin); Golfo de Uraba, CNHM 63881 (Goin); Pena Lisa, Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro, CNHM 4451-3, 4455-61 (Goin); Rio San Juan, Playa del Oro, CNHM 54772 (Goin); Rio Quesada, AMNH 13615-77; 37 km. up Rio Pune, AMNH 13688; 48 km. up Rio Pune, AMNH 13689. =Narino=: Tumaco, Rio Rosario, CJG 2310-13 (Goin). =Valle=: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT 166, 346-47, 388 (Goin); Rio Calima above Cordoba, CJG 2249-57 (Goin).

ECUADOR: No province: Bulun, AMNH 10620. =Esmeraldas=: Cachabe, AMNH 10625-8; Rio Capayas, CNHM 35712; Rio Sapaya, UMMZ 58910 (5); Salidero, AMNH 10623-4; San Javier, AMNH 10618. =Guayas=: Hacienda Balao Chico, UMMZ 123904. =Imbabura=: Pambelar, AMNH 10629, 10631. =Pichincha=: Hacienda Espinosa, 9 km. W Santo Domingo de los Colorados, KU 40220.

=Smilisca puma= (Cope), new combination

_Hyla puma_ Cope, Proc. Amer. Philos. Soc., 22:183, 1885 [Holotype.--USNM 13735 from Nicaragua; J. F. Moser collector].

Gunther, Biologia Centrali-Americana: Reptilia and Batrachia, p.

270, Sept., 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 251, June, 1923. Cochran, Bull. U. S. Natl. Mus., 220:58, 1961.

_Hyla wellmanorum_ Taylor, Univ. Kansas Sci. Bull. 25(1):843, July 1, 1952 [Holotype.--KU 30302 from Batan, Limon, Costa Rica, Edward H. Taylor collector]; Univ. Kansas Sci. Bull., 36(1):626, June 1, 1954. Duellman and Berg, Univ. Kansas Publ. Mus. Nat. Hist., 15:194, Oct. 26, 1962.

_Smilisca wellmanorum_, Starrett, Copeia, 4:303, Dec. 30, 1960.

_Diagnosis._--Size small ([M] 38.0 mm., [F] 46.0 mm.), differing from other species in the genus by the following combination of characters: skull about as long as broad; frontoparietal fontanelle keyhole-shaped; supraorbital f.l.a.n.g.es absent; squamosal small, not in contact with maxillary; bony portion of ethmoid terminating at anterior edge of orbit; tarsal fold weak, two-thirds length of tarsus; inner metatarsal tubercle small, low, flat, elliptical; snout rounded in dorsal profile; lips thin and flaring; fingers having only vestige of web; toes one-half webbed; diameter of tympanum about two-thirds that of eye; narrow l.a.b.i.al stripe white; pair of dark brown (sometimes interconnected) stripes on tan dorsum; no blue spots on flanks or thighs; vocal sac in breeding males pale brown. (Foregoing combination of characters distinguis.h.i.+ng _S. puma_ from other species in genus.)

_Description and variation._--Ten breeding males from the vicinity of Puerto Viejo, Heredia Province, Costa Rica, have snout-vent lengths of 32.5 to 37.9 mm. (34.8 mm.). In these specimens, the length of the tibia to the snout-vent length is 0.48 to 0.53 (0.51), and the tympanum/eye ratio is 0.52 to 0.72 (0.65). Seven females have snout-vent lengths of 40.8 to 45.8 mm. (43.9 mm.). No individual has more than a vestige of a web between the second and third and fourth fingers. None has a web between the first and second fingers. Breeding males lack nuptial excrescences on the thumbs. The vocal sac is moderately large and bilobed.

In preserved specimens the dorsal ground color varies from yellowish tan to grayish brown. All specimens have dark brown dorsal markings in the form of a pair of dorsal stripes, variously modified (Pl. 7A). In some specimens, such as KU 91716, the stripes are discrete and extend from the pos...o...b..tal region nearly to the vent. In most specimens the stripes are connected by a transverse mark in the scapular region and in many others also by a transverse mark in the sacral region. In some specimens the stripes are fragmented posteriorly; fragmentation is extreme in KU 30300, in which the dorsal pattern consists of two series of dark longitudinal dashes. The other extreme is a nearly complete fusion of the stripes, as in KU 91714. A dark brown interorbital bar usually extends onto the eyelids, but in some specimens this is reduced to a short V-shaped mark or small spot between the eyes. There is no dark post-tympanic mark, but dark brown pigment forms a venated pattern from the axilla to the mid-flank; the inguinal region is white, finely mottled with dark brown. The dorsal surfaces of the hind limbs are colored like the body and have two or three dark brown transverse marks on the thighs, three to five marks on the shanks, and one or two marks or irregularly arranged dark flecks on the tarsi. The anterior and posterior surfaces of the thighs are pale tan to brown. The webbing of the feet is tan to grayish brown. A narrow white l.a.b.i.al stripe, white a.n.a.l stripe, and narrow white stripes on the tarsi and outer edges of the forelimbs are invariably present. The ventral surfaces are creamy white.

In life the dorsum is tan or pale brown with dark brown markings. Some individuals have scattered metallic green flecks on the dorsum. The flanks are mottled dark brown and creamy white. The posterior surfaces of the thighs are dark brown. The vocal sacs are grayish brown, and the iris is a deep bronze color.

_Natural History._--_Smilisca puma_ inhabits humid lowland tropical forests having more or less evenly distributed rainfall throughout the year. The equable climatic conditions seemingly permit these frogs to be active throughout most of the year. Taylor (1952:846) found calling males at Batan, Costa Rica, on July 20, 1951. We found the species breeding near Puerto Viejo, Costa Rica, on February 19, June 18, July 13, and July 31. Specimens of calling males from Costa Rica in the collection at the University of Southern California were obtained in February at La Fortuna, on August 22 at Los Diamantes, on August 30 at Jabillos, and on September 5 at La Lola. Gravid females were collected in June, July and August.

Males call from shallow water. All breeding congregations of this species that we have found were in a gra.s.sy marsh, 7.5 kilometers west of Puerto Viejo, Costa Rica. Tadpoles were found in water-filled depressions in the marsh at night. When first observed, tadpoles were near the surface of the water; they responded to light by quickly taking refuge in the dense gra.s.s. No tadpoles were observed by day.

The breeding call consists of a low squawk, usually followed by a series of one or more rattling secondary notes (duration of primary notes, 0.06-0.35 seconds; of secondary notes, 0.10 to 0.47 seconds), repeated at intervals of 5 to 55 seconds. The primary notes have 187 to 240 pulses per second and major frequencies of about 740 to 1870 cycles per second (Pl. 11A).

Only six tadpoles are available for study. Four of them in stage 34 of development have body lengths of 9.0 to 9.5 mm., tail lengths of 14.0 to 15.0 mm., and total lengths of 23.0 to 24.5 mm. One tadpole in stage 38 and one in stage 40 have total lengths of 31.0 mm. A typical tadpole in stage 34 of development (KU 91807 from 7.5 km. W Puerto Viejo, Heredia Province, Costa Rica) has a body length of 9.5 mm., tail length of 15.0 mm., and total length of 24.5 mm.; body about three-fourths as deep as wide; snout rounded dorsally and laterally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, about two-thirds distance from snout to posterior end of body and slightly below midline; a.n.a.l tube dextral; caudal musculature slender, barely curved upward distally; dorsal fin extending onto body; at mid-length of tail, depth of caudal musculature equal to that of dorsal fin and ventral fin; body grayish brown, palest ventrally; caudal musculature pale creamy yellow with bold gray reticulations; caudal fins transparent with gray reticulations anteriorly and black flecks posteriorly on both fins (Fig.

14A). Median part of upper lip bare; rest of mouth bordered by two rows of short blunt papillae; lateral fold present; tooth-rows 2/3; upper rows equal in length; second upper row broadly interrupted medially; three lower rows complete, first and second rows equal in length, slightly shorter than upper rows; third lower row noticeably shorter; upper beak shallow, forming broad, continuous arch with slender lateral processes; lower beak slender, broadly V-shaped, both beaks finely serrate (Fig. 15B).

All six tadpoles are colored alike, except that in the larger specimens scattered white flecks are present on the ventral surface of the body, and the dark reticulations continue farther posteriorly on the caudal fins than in the smaller tadpoles. In two specimens the third lower tooth-row is only about one-half the length of the other lower rows, and in one specimen the second lower tooth-row is shorter than the first.

Coloration of tadpoles in life: "Body olive-brown with silvery green flecks laterally. Caudal musculature olive-brown with greenish tan flecks. Fins brown with greenish gold flecks. Iris deep bronze."

(Duellman, field notes, February 19, 1965).

One recently metamorphosed young (KU 91808) has a snout-vent length of 12.4 mm. In life this frog had a pale tan dorsum with dark brown markings, yellowish tan posterior surfaces of thighs, grayish brown throat, and bronze iris.

_Remarks._--The ident.i.ty of Cope's _Hyla puma_ has not been known. The name has appeared in various compilations, but no workers have referred any of their specimens to that species. Examination of the holotype (USNM 13735), an adult female, revealed the presence of the following combination of characters: snout-vent length 45.8 mm., snout blunt above and rounded laterally, nostrils close to tip of snout, lips thin and flaring, a vestige of a web on the hands, feet about one-half webbed, tarsal fold weak and extending about two-thirds length of tarsus, dorsal markings consisting of a faded dark interorbital bar and a pair of faded longitudinal brown marks connected by a transverse band in the scapular region. The type agrees well with specimens of _Smilisca wellmanorum_ (Taylor, 1952); the vestigial webbing on the hands and the dorsal coloration are especially significant. Consequently, we consider _Hyla wellmanorum_ Taylor, 1952, to be a synonym of _Hyla puma_ Cope, 1885.

Cope gave only "Nicaragua" as the locality for _Hyla puma_. The specimen was part of a collection received at the United States National Museum from Lt. J. F. Moser. Among the species in the collection are _Dentrobates pumilio_, _Phyllomedusa helenae_, _Corythophanes cristatus_, _Pliocercus dimidatus_, _Tretanorhinus nigroluteus_, and others characteristically found on the Caribbean lowlands of Central America. Thus, it seems reasonable to a.s.sume that the type specimen of _Hyla puma_ came from the Caribbean lowlands. Though no other Nicaraguan specimens have been found by us, numerous specimens are known from the Caribbean lowlands of Costa Rica.

Cochran (1961:58), in her catalogue of type specimens in the United States National Museum, listed _Hyla puma_ Cope, 1885, as a synonym of _Hyla molitor_ Schmidt, 1857. She made no qualifying statements. Schmidt (1858:246), in his descriptions of the species in the year following his publication of the names and Latin diagnoses, stated: "Dorsum uniformly gray, more intensive on back, fading away laterally and on extremities; in every-day-life this blue would be called _Mueller's Blau_. A delicately dotted black line runs on the canthus rostralis from the opening of the nose to the corner of the eye. In the armpits, on the flanks and the thighs two of our three specimens have black marblings."

[Free translation] Certainly on the basis of coloration _Hyla puma_ is distinctly different from _Hyla molitor_.

_Distribution._--This species lives in the wet, forested regions of the Caribbean lowlands of Costa Rica and presumably southern Nicaragua (Fig.

3). All specimens are from low elevations; the highest known elevation for the occurrence of this frog is 285 meters at Laguna Bonilla.

[Ill.u.s.tration: FIG. 3. Map showing locality records for _Smilisca puma_ (triangles) and _Smilisca sila_ (circles).]

_Specimens examined._--62, as follows: NICARAGUA: No specific locality, USNM 13735.

COSTA RICA: =Alajuela=: Jabillos, 5 km. N Santa Clara, USC 8058 (6); 5 km. W La Fortuna, USC 8078 (2); Rio La Fortuna at La Fortuna, USC 7151 (3). =Cartago=: Laguna Bonilla, tunnel camp near Peralta, KU 32171.

=Heredia=: Puerto Viejo, KU 86521; 5.9 km. W Puerto Viejo, KU 64307; 7.5 km. W Puerto Viejo, KU 64308-10, 64311 (skeleton), 64312-15, 68635-6 (skeletons), 85001-2 (skeletons), 86520, 87770-1 (skeletons), 91709-16, 91791-2, 91807 (tadpoles), 91808 (young). =Limon=: Batan, KU 30300-2; La Lola, KU 32169, USC 141, 201, 8067; Los Diamantes, KU 32170, UMMZ 118470 (6), USC 212; 2.4 km. E Los Diamantes, USC 8049 (5).

=Smilisca sila= new species

_Hyla gabbi_, n.o.ble, Proc. Biol. Soc. Was.h.i.+ngton, 37:66, Feb. 21, 1924. Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, Oct. 10, 1931. Schmidt, Smithsonian Misc. Coll., 89(1):6, March 16, 1933.

_Hyla sordida_, Dunn, Copeia, 3:166, Nov. 19, 1937. Cooper, Copeia, 2:121, June 30, 1944. Breder, Bull. Amer. Mus. Nat. Hist., 86(8):417, Aug. 26, 1946.

_Hyla phaeota_, Breder, Bull. Amer. Mus. Nat. Hist., 86(8): pl. 55, Aug. 26, 1946.

_Holotype._--Adult male, KU 91852 from a small stream at the north edge of the village of El Volcan, Chiriqui Province, Panama, elevation 1280 meters; obtained on Feb. 5, 1965, by William E. Duellman.

_Paratypes._--KU 91853-74, collected with the holotype.

_Diagnosis._--Size moderate ([M] 45.0 mm., [F] 62.2 mm.); skull wider than long, having large, ovoid frontoparietal fontanelle; supraorbital f.l.a.n.g.es absent; squamosal small, not contacting maxillary; bony section of ethmoid extending anteriorly between nasals; tarsal fold weak, full length of tarsus; inner metatarsal tubercle low, flat, elliptical; lips thick, rounded, not flaring; fingers one-third webbed; toes three-fourths webbed; diameter of tympanum about one-half that of eye; margin of upper lip faintly marked by interrupted creamy white stripe; dark spots on dorsum; pale flecks on flanks and posterior surfaces of thighs; vocal sacs in breeding males dark brown. (Foregoing combination of characters distinguis.h.i.+ng _S. sila_ from any other species in genus.)

_Description of holotype._--Snout-vent length 36.6 mm.; tibia length 19.8 mm., 54.1 per cent of snout-vent length; foot length 15.5 mm., 42.3 per cent of snout-vent length; head length 12.7 mm., 34.7 per cent of snout-vent length; head width 13.3 mm., 36.8 per cent of snout-vent length; snout short, in lateral profile truncate, only slightly rounded above, in dorsal profile rounded; canthus rounded; loreal region noticeably concave; lips thick, rounded, not flaring; nostrils not protuberant, directed laterally; internarial distance 3.0 mm.; internarial area flat; top of head flat; interorbital distance 3.5 mm., 26.3 per cent of head width; diameter of eye 4.2 mm., thrice distance (1.4 mm.) from tympanum to eye, and half again distance (2.8 mm.) from orbit to nostril; pupil horizontally ovoid; width of eyelid 2.8 mm., 21.1 per cent of head width; dermal fold from posterior corner of orbit covering upper edge of tympanum to point above insertion of forelimb; diameter of tympanum 2.3 mm., 54.7 per cent of diameter of eye; no axillary membrane; arms moderately robust; weak fold on wrist; faintly scalloped fold along ventrolateral margin of forearm; fingers short, slender; fingers from shortest to longest, 1-2-4-3; vestige of web between first and second fingers; others about two-fifths webbed; discs moderate, diameter of that on third finger about one-third diameter of eye; triangular outer palmar tubercle; elliptical inner palmar tubercle on base of pollex; subarticular tubercles large, conical, none bifid; supernumerary tubercles few, large, conical; brown nuptial excrescence on prepollex; heels overlap by about one-fifth length of shank when hind limbs adpressed; tibiotarsal articulation extending to nostril; tarsal fold weak, extending nearly full length of tarsus; inner metatarsal tubercle elliptical, flat; outer metatarsal tubercle absent; toes moderately long; toes from shortest to longest, 1-2-3-5-4, third and fifth about equal in length; discs about same size as those on fingers; webbing extending to middle of penultimate phalanx on all toes, except only to distal end of antepenultimate phalanx of fourth toe; subarticular tubercles round; supernumerary tubercles large, round, present only on proximal digits. a.n.a.l opening directed posteriorly at level of upper edge of thighs; no noticeable a.n.a.l sheath; flat tubercles ventrolateral to a.n.a.l opening large; skin of chest, belly, and posterior surfaces of thighs granular; other surfaces smooth; tongue broadly cordiform, shallowly notched posteriorly, and barely free behind; vomerine teeth 4-4, situated on ventral surfaces of separated rounded prominences between posterior margins of small, ovoid inner nares; vocal slits long, each situated along inner margin of ramus; color (in preservative) pinkish tan above with irregular olive-brown markings forming interconnected spots on back; four bars on dorsal surface of each thigh; five bars on shank, and three on tarsus; inguinal region white with black mottling; posterior surfaces of thighs yellowish tan proximally, dark brown distally; margins of lips grayish white with brown markings; ventral surfaces of hands and feet grayish brown; belly and posterior part of throat creamy white; anterior part of throat brown.

_Description and variation._--Ten breeding males from Finca La Sumbadora, Panama, have snout-vent lengths of 40.0 to 44.8 mm. (42.3 mm.). In these specimens the tibia/snout-vent length ratio is 0.50 to 0.57 (0.54), and the tympanum/eye ratio is 0.48 to 0.58 (0.53). There is a geographic gradient in size; specimens from the western part of the range (southern Costa Rica) are smaller than those from the eastern part of the range (eastern Panama). Five males from the Pacific lowlands of southern Costa Rica have snout-vent lengths of 31.6 to 38.2 mm. (34.7 mm.); ten males from El Volcan, Chiriqui, Panama, 32.6 to 37.9 mm. (36.4 mm.), and eight males from Barro Colorado Island, Ca.n.a.l Zone, 38.2 to 42.0 mm. (35.6 mm.). These are smaller than the males from Finca La Sumbadora, which is east of the Ca.n.a.l Zone. Ten females from El Volcan have snout-vent lengths of 44.2 to 55.6 mm. (49.2 mm.), as compared 56.1 to 62.2 mm. (58.2 mm.) in three females from Finca La Sumbadora.

Large females have scattered small tubercles on the head and back; tubercles occur in males from Costa Rica and in some males from western Panama. The truncate snout is characteristic of both s.e.xes.

The coloration of _Smilisca sila_ consists of a gray, tan, or pale reddish brown dorsal ground color and a creamy white venter. The dorsum is marked by dark brown, olive-brown, or dark reddish brown spots or blotches (Pl. 7B). Usually the blotches are discrete, but in some individuals they are interconnected and form an irregular dark mark on the dorsum. There is no tendency for the blotches to form transverse bars as in _Smilisca sordida_. In one specimen (KU 80467) the blotches are fused and form two wide irregular longitudinal stripes, as in _Smilisca puma_. In some females the dorsal markings are reduced to a few small spots or are nearly absent (KU 92332), whereas in other females the dorsal markings are bold. In one female (KU 91894) the dorsal markings are narrowly bordered by pale blue, and numerous pale blue flecks are present on the pale brown dorsum. In many individuals of both s.e.xes small white flecks are present on the dorsal surfaces.

Usually the flanks and posterior surfaces of the thighs have black mottling enclosing pale blue spots and flecks, respectively. The dorsal surfaces of the limbs are marked by dark brown transverse bars; usually three or four bars are present on each forearm, thigh, and shank. The coloration of the flanks and limbs varies geographically. Specimens from southern Costa Rica and western Panama have distinct bars on the limbs; the posterior surfaces of the thighs have brown reticulations enclosing small blue flecks in specimens from Costa Rica and bolder, black reticulations enclosing large pale blue spots in specimens from western Panama. In specimens from Costa Rica the flanks are brown with pale blue flecks, whereas in those from Chiriqui, Panama, the flanks are pale blue with dark brown mottling in the inguinal region. Frogs from El Valle and Cerro la Campana usually have distinct bars on the limbs; the posterior surfaces of the thighs are colored as in frogs from Chiriqui, and the inguinal region is pale blue with coa.r.s.e brown mottling. Specimens from Barro Colorado Island are marked like those from El Valle and Cerro la Campana, except that on the posterior surfaces of the thighs fine black reticulations enclose many dark blue spots. In specimens from Darien and from Panama Province east of the Ca.n.a.l Zone (Altos de Pacora, Cerro Jefe, Finca La Sumbadora, and Rio Pacora), the markings on the dorsal surfaces of the limbs are indistinct or absent in males, but distinct in some females. Intense brown and black pigment forms fine reticulations delimiting bold blue spots on the flanks; this coloration extends to the axilla in many specimens. Fine black reticulations enclose many dark blue spots on the posterior surfaces of the thighs.

In females, the throat is creamy white; in some specimens scattered brown flecks are present on the chin and throat. In breeding males the anterior part of the throat is dark gray or dark brown.

The coloration in life is as variable as it is in preservative. In life the holotype had a tan dorsum with dark olive-green irregular markings and small green flecks. The limbs were tan with dark brown transverse bars. The flanks were grayish tan anteriorly; the inguinal region and posterior surfaces of thighs were blue with black mottling. The belly was creamy white, and the throat was brown with creamy yellow flecks.

The iris was a dull bronze color. Among the paratypes, some individuals had green flecks, others did not. The inguinal region and posterior surfaces of the thighs were pale blue, pale yellowish green, or grayish tan with black mottling. The blue was most noticeable in females.

Colors of a male from Finca La Sumbadora, Panama, were described as follows: "Dorsum olive-brown; irregular dark brown blotches, pale green flecks, and raised creamy yellow spots on dorsal surfaces; belly creamy white; throat grayish brown; undersides of limbs grayish tan; groin, anterior and posterior surface of thigh, inner surface of shank, anterior edge of tarsus, and proximal parts of third and fourth toes pale blue marbled with dark brown and black; webbing brown; iris pale bronze, finely reticulated with black." (Duellman, field notes, January 28, 1964.)