Part 9

Smith (1953:150) described Hyla phaeota cyanosticta from Guatemala. Our studies on the external morphology, coloration, and especially the cranial osteology provide evidence that cyanosticta is a species distinct from phaeota.

Distribution.--Smilisca phaeota inhabits humid tropical forests from northeastern Nicaragua southward on the Caribbean lowlands to elevations of about 1000 meters and on the Pacific lowlands of Costa Rica, exclusive of the arid regions of Guanacaste, throughout the lowlands of Panama, exclusive of the savannas of the Pacific lowland and the Azuero Peninsula, and southward on the Pacific slopes of South America through Colombia to west-central Ecuador; also the valleys of the Rio Cauca and Rio Magdalena in Colombia (Fig. 2).

Specimens examined.--528, as follows: NICARAGUA: =Matagalpa=: Finca Tepeyac, 10 km. N, 9 km. E Matagalpa, KU 85439, 87707 (tadpoles); Matagalpa, MCZ 3546-7, UMMZ 92367; 19 km. N Matagalpa, UMMZ 116495-6. Zelaya: Bonanza, KU 84854-62, 84950-2 (skeletons), 85440-50, 87708-9 (tadpoles); Cukra, AMNH 80618; Rio Mico, 16 km. E Recreo, UMMZ 79711 (6), 79712 (4); junction Rio Mico and Rio Siguia, UMMZ 79713 (10); Rio Siguia, 11 km. NW Rama, UMMZ 79714 (14), 79715 (11), 79716 (21), 79717, 79718 (3).

COSTA RICA: =Alajuela=: Cinchona, KU 32255, 64286-8; 5 km. S Ciudad Quesada, USC 8077; Laguna Monte Alegre, KU 64289-90; Las Playuelas, 11 km. S Los Chiles, USC 7216; San Carlos, USNM 29961.

=Cartago=: Moravia de Turrialba, KU 32212-47, 37133-5, 41093 (skeleton), 64280-1, USC 7243 (3); Peralta, KU 32271-2; Rio Chitaria, 3 km. NNE Pavones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); Rio Reventazon, MCZ 29196-203, UMMZ 117677 (9); Turrialba, KU 25720-2, 32209-11, 32266-8, 32273-4, 37136-67, 41090-2 (skeletons), 64270-2, MCZ 29221, 29222 (tadpoles), 29269-70, USNM 29934.

=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. =Canal 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]. Guenther, 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 ( 38.0 mm., 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 flanges 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 labial 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 distinguishing 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 postorbital 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 labial stripe, white anal 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 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.