Males call from shallow water. All breeding congregations of this species that we have found were in a grassy 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 grass. 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; anal 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 identity 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 assume 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.