Part 10

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.

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, Noble, Proc. Biol. Soc. Washington, 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 ( 45.0 mm., 62.2 mm.); skull wider than long, having large, ovoid frontoparietal fontanelle; supraorbital flanges 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 distinguishing 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. Anal opening directed posteriorly at level of upper edge of thighs; no noticeable anal sheath; flat tubercles ventrolateral to anal 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, Canal 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 Canal 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 sexes.

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 sexes 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 coarse 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 Canal 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.)

A female (now KU 91894) from Altos de Pacora, Panama, was described as follows: "An irregular dark brown, green-bordered figure on head and back; dark brown, green-bordered bands on limbs--all on a lighter brown and heavily green-spotted background. These markings are more vivid at night than during the day. Lower sides, from midbody onto front of thighs and rear of thighs onto venter of shanks to heels and thence dorsally onto basal portions of toes heavily blue spotted on a light brown (front of thighs and venter of shanks) to blackish brown background. Venter cream. Iris gray-brown, finely veined with dark brown." (Charles W. Myers, field notes, December 14, 1964.) Note that in the earlier discussion of coloration of preserved specimens, the green spots and borders have changed to pale blue after six months in alcohol.

In living individuals from Costa Rica and Panama west of the Canal Zone, the blue coloration on the flanks and thighs is much less conspicuous than in specimens from eastern Panama. The color of the iris is variable, even in frogs from one locality. The coloration of the iris in 13 living frogs (now KU 92333-45) from Valle Hornito, Chiriqui, Panama, was described as follows: "Iris variable--from pale to dark brown; in a few the iris has a golden cast to the brown; in a few others the lower half of the iris is pale gray with the upper half being light brown." (Charles W. Myers, field notes, April 24, 1965).

Natural history.--Smilisca sila inhabits the Pacific slopes of lower Central America where a pronounced dry season occurs. We have records of males calling in December through May and also in August (latter date from El Volcan, Chiriqui, Panama). The breeding season seems to be correlated with the time of the year when the water is clear and at a low level in the streams where these frogs breed.

Males call from the edges of small, shallow streams, from rocks in the streams, or less frequently from vegetation overhanging the streams. Females are most frequently found on the banks of streams, and clasping pairs usually are in shallow pools in streams. One individual was found in a bromeliad about three meters above the ground in the daytime.

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 to 0.28 seconds; of secondary notes, 0.14 to 0.48 seconds), repeated at intervals of 4 to 20 seconds. The primary notes have 97 to 120 pulses per second and major frequencies of about 900 to 2220 cycles per second (Pl. 11B).

Eggs were obtained artificially in the field; the average length of ten embryos in the neural groove stage is 2.4 mm., and the average diameter of the outer envelope is 4.9 mm. Hatchlings have large, conical oral discs, heavy gills, and a large amount of yolk; their average total length is 6.3 mm.

Tadpoles have been found in pools in clear streams; some tadpoles have been observed to cling by their mouths to rocks in the stream; others were found on the bottom where they seek refuge among pebbles or under rocks and leaves. A complete developmental series of tadpoles is not available. Eleven tadpoles in stage 25 of development have body lengths of 8.3 to 10.2 mm. (9.3 mm.), tail lengths of 17.3 to 21.0 mm. (18.8 mm.), and total lengths of 25.9 to 31.0 mm. (28.1 mm.). One tadpole in stage 41 and one in stage 42 have body lengths of 11.5 and 12.5 mm., tail lengths of 27.2 and 29.5 mm., and total lengths of 38.7 and 42.0 mm., respectively. The snout-vent lengths of two specimens in stage 43 and one in stage 45 are 12.7, 13.0, and 13.6 mm., respectively.

A typical tadpole in stage 25 of development (KU 80620 from Finca La Sumbadora, Panama) has a body length of 9.5 mm., tail length of 19.0 mm., and a total length of 28.5 mm.; body only slightly wider than deep, nearly flat dorsally; snout broadly rounded in dorsal view, bluntly rounded in lateral view; eyes widely separated, directed dorsolaterally; nostril slightly closer to eye than to tip of snout; mouth ventral; spiracle sinistral, located about two-thirds distance from snout to posterior edge of body; anal tube dextral; caudal musculature moderately heavy, straight; dorsal fin not extending onto body; fins deepest at about two-fifths length of tail, where depth of caudal musculature about equal to depth of dorsal and depth of ventral fin; musculature extending nearly to tip of tail; body dark grayish brown above and pale grayish tan below with small dark brown spots dorsally and white flecks laterally; caudal musculature pale tan with dark brown flecks over entire surface and dark brown streaks on posterior one-half of ventral fin and on all of dorsal fin (Fig. 14B). Median one-third of upper lip bare; rest of mouth bordered by a single row of conical papillae; lateral fold present; tooth rows 2/3; upper rows cone-shaped, about equal in length, broadly /\-shaped; second upper row narrowly interrupted medially; lower rows complete, about equal in length, but slightly shorter than upper rows; upper beak moderately massive, its inner surface forming a continuous arch with short lateral processes; lower beak broadly \/-shaped; both beaks finely serrate (Fig. 15D).

Tadpoles from El Volcan, Chiriqui (KU 91833), are more heavily pigmented than those from Finca La Sombadora; the spots on the tail are larger. In life these tadpoles had dark brownish black bodies with golden and green lichenous flecks; the tail was tan with dark brown markings, and the iris was a grayish bronze color. In life tadpoles from Finca La Sumbadora were olive-tan above and dark gray with pale bluish gray irridescent spots ventrally. The caudal musculature was creamy tan with brown flecks and streaks, and the iris was pale bronze.