Specimens examined.—78, as follows:Mexico:Chiapas: Monte Libano, MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH 41588.Oaxaca: 11 km. N Vista Hermosa, KU 84918-20 (skeletons), 87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH 57199-201; 8 km. S Yetla, KU 87213, UMMZ 124838 (8).Veracruz: Coyame, UMMZ 111459-60; between Coyame and Tebanco, UMMZ 121198; Dos Amates, UMMZ 121297; between Laguna de Catemaco and Volcán San Martín, UMMZ 121200; Volcán San Martín, UIMNH 35403-4, 35408-12, UMMZ 118163; SE slope Volcán San Martín, UMMZ 121199, 121295 (2), 121296, 121298.Guatemala:Alta Verapaz: Chinajá, KU 55935-7, 55938 (skeleton).ElPetén: 10 km. NNW Chinajá (Alta Verapaz), KU 55934; Piedras Negras, CNHM 99006-7, 99227, UIMNH 28853, USNM 111139-41, 111143-7; 8 km. S Piedras Negras, CNHM 99008; Semicoch, USNM 35907.Fig. 2.Map showing locality records forSmilisca cyanosticta(triangles) andSmilisca phaeota(circles).Smilisca phaeota(Cope)Hyla phaeotaCope, Proc. Acad. Nat. Sci. Philadelphia, 14 (9):358, 1862 [Holotype.—USNM 4347 from Turbo, Colombia; J. Cassin collector]. Boulenger, Catalogue Batrachia Salientia in British Museum, p. 402, Feb. 1, 1882. Werner, Sitzungsb. Akad. Wiss. München, 27:215, 1897. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p. 269, Sept. 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 261, June 1923. Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, Oct. 10, 1931. Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 357:4, Oct. 26, 1937. Cooper, Copeia, 2:122, June 30, 1944. Breder, Bull. Amer. Mus. Nat. Hist., 86(6):416, Aug. 26, 1946. Smith and Taylor, Bull. U.S. Natl. Mus., 194:88, June 17, 1948; Univ. Kansas Sci. Bull, 33:364, March 20, 1950. Taylor, Univ. Kansas Sci. Bull., 35(1):837, July 1, 1952. Brattstrom and Howell, Herpetologica, 10:117,[Pg 309]Aug. 1, 1954. Goin, Herpetologica, 14:120, July 23, 1958. Cochran, Bull. U.S. Natl. Mus., 220:57, 1961.Hyla labialisPeters, Monats. Konigl. Akad. Wissen. Berlin, p. 463, 1863 [Holotype.—ZMB 4913 from "region of Bogotá," Colombia]; Monats. Konigl. Akad. Wissen. Berlin, p. 618, Oct. 16, 1873. Boulenger, Catalogue Batrachia and Salientia in British Museum, p. 397, Feb. 1, 1882.Hyla baudini dolomedesBarbour, Occas. Papers Mus. Zool. Univ. Michigan, 129:11, Jan. 25, 1923 [Holotype.—MCZ 8539 from Río Esnápe, Sambú Valley, Darién, Panamá; Barbour and Brooks collectors]. Barbour and Loveridge, Bull. Mus. Comp. Zool. Harvard, 69:278, June, 1929.Hyla phaeota phaeota, Smith, Herpetologica, 8:152, Jan. 30, 1953. Minton and Smith, Herpetologica, 16:103, June 17, 1960.Smilisca phaeota, Starrett, Copeia, 4:303, Dec. 30, 1960.Diagnosis.—Size large ([M] 65 mm., [F] 78 mm.); skull as long as wide, lacking frontoparietal fontanelle; large supraorbital flanges having straight edges and extending posterolaterally; large squamosal not in contact with maxillary; tarsal fold moderately wide, full length of tarsus; inner metatarsal tubercle moderately large, low, flat, elliptical; hind limbs relatively long, tibia averaging more than 53 per cent of snout-vent length; labial stripe silvery white; lips lacking vertical bars; loreal region pale green; dark brown or black tympanic mark dispersing into brown venated pattern on flanks; posterior surfaces of thighs pale brown, with or without darker flecks or small cream-colored flecks. (Foregoing combination of characters distinguishingS. phaeotafrom any other species in genus.)Table 2.—Geographic Variation in Size and Proportions in Males of Smilisca phaeota. (Means in Parentheses Below Observed Ranges; Data Based of Ten Specimens From Each Locality.)LocalitySnout-ventlengthHead width/snout-vent lengthInterorbital distance/head widthBonanza, Nicaragua40.8-47.734.1-38.031.0-36.1(43.7)(36.3)(35.4)Heredia Prov., Costa Rica46.3-59.032.5-36.030.5-39.6(51.7)(35.0)(34.7)Puntarenas Prov., Costa Rica53.6-64.932.6-36.131.0-38.0(61.4)(34.5)(34.4)Canal Zone, Panamá52.4-65.533.5-37.631.3-37.2(56.5)(35.6)(34.7)Río Quesada, Colombia52.6-61.033.1-37.130.1-33.9(56.0)(35.0)(32.1)Description and Variation.—For the purposes of analyzing geographic variation in size and proportions, measurements were taken on ten adult males from each of five samples throughout the range of the species. Aside from the data summarized in Table 2, the average ratio of tibia length to snout-vent length is noticeably less in Colombian specimens (53.4 per cent, as compared with 54.8 to 57.8 per cent in the other samples) and the ratio of head length tosnout-vent length is noticeably less in Costa Rican specimens (33.5 per cent as compared with 34.9 to 35.1 per cent in the other samples). Also, specimens from Heredia Province, Costa Rica, have a relatively smaller tympanum (62.7 to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared with means of 74.0 to 77.9 per cent in the other samples).Two populations are distinctive as regards the size of adult males. Specimens from the northern Caribbean lowlands of Nicaragua (Bonanza, the northernmost locality for the species) are remarkably small. Males having snout-vent lengths of between 40 and 43 mm. were breeding; the largest male found had a snout-vent length of 47.7 mm. The other extreme in size is attained in specimens from the Pacific lowlands of eastern Costa Rica and western Panamá, where most breeding males have snout-vent lengths of more than 55 mm.; the largest male had a snout-vent length of 64.9 mm.The rather striking differences in size among certain samples and the minor differences in proportions among other samples show no geographic trends. Instead, the variations apparently are random among the samples. The data presented here possibly are the results of inadequate sampling, but more likely reflect actual differences in the populations.The dorsal ground color ofSmilisca phaeotais pale green to tan; the venter is creamy white. The dorsum is variously marked with dark olive-green or dark brown spots or blotches (Pl. 6C). A dark interorbital bar is usually present. Usually a large dark dorsal mark extends from the occiput to the sacral region, but in many individuals this blotch is replaced by two or three dark marks. The dorsal markings are irregular in shape and do not tend to form transverse bands or longitudinal bars. The hind limbs are marked by dark transverse bands, usually four or five on the thigh, five or six on the shank, and four on the tarsus. Two or three narrow bands are usually present on the proximal part of the fourth toe. The webbing on the feet is brown. The loreal region is pale green, bordered above by a narrow dark brown canthal stripe extending from the nostril to the orbit. The upper lip is silvery white. A broad dark brown or black mark extends posteriorly from the orbit, encompassing the tympanum, to a point above the insertion of the forelimb. The flanks are pale green or pale tan and marked with a fine dark brown or black venation. The anterior surfaces of the thighs usually are pale brown or grayish tan, sometimes having small, indistinct darker flecks. The posterior surfaces of the thighs are similarly colored, but in most specimens small but distinct dark flecks are present; in some specimens small cream-colored spots are also present on the posterior surfaces of the thighs. A distinct, narrow creamy white anal stripe usually is present. A distinct white stripe is present on the outer edge of the tarsus and fifth toe; on the tarsus the white stripe is bordered below by dark brown. A white stripe also is present on the outer edge of the forearm and fourth finger. In breeding males the throat is dark gray.Little geographic variation in color or pattern is evident. Few, if any, specimens from the Pacific lowlands of South America are green in life. (We have seen no living individuals from South America.) Some living individuals from Costa Rica and all those seen alive from Nicaragua have a tint of pale blue on the flanks. In some specimens the dorsal pattern is so faint as to be barely discernible, whereas in most specimens the pattern is bold.The coloration in the living frogs is highly variable due to extreme metachrosis. Individuals of this species are capable of changing the dorsal colorationfrom green to brown in a short period of time. Both green and brown individuals have been found active at night. Usually those individuals found hiding by day are brown. One individual from Finca La Sumbadora, Panamá (now KU 91914), was kept alive in the laboratory for nearly one month. This individual usually was pale green with tan dorsal markings at night and tan with pale green markings by day. On occasion the pale green dorsal markings were boldly outlined by bright dark green.In living individuals from throughout the range of the species the iris is a bronze color, darkest medially with fine black reticulations.Natural History.—Smilisca phaeotainhabits humid lowland tropical forest and seldom ascends the foothills to more than 1,000 meters. The rather equable climatic conditions, especially more or less evenly distributed rainfall throughout the year, permit this frog to be active most of the year. Dunn (1931:413) reported males calling on Barro Colorado Island, Panamá, in February and in July, and Breder (1946:416) noted calling individuals in the Chucanaque drainage of Darién, Panamá in January, March, July, August and October and in Costa Rica in April through August inclusively. Calling males were found at Bonanza, Nicaragua in March and in July.At all times of year the usual daytime retreats for these frogs are near water; the frogs have been found in elephant ear plants (Xanthosoma) and in bromeliads; occasional individuals have been found sitting on shaded branches of bushes and trees. None has been observed on the ground or beneath ground-cover by day.The length of the breeding season cannot be determined definitely. The earliest date on which eggs have been found is May 23; Gaige, Hartweg, and Stuart (1937:5) reported a gravid female taken at El Recreo, Nicaragua, in September, and we have a gravid female taken at Almirante, Panamá, in March.Males usually call from secluded spots at the edge of water. All calling males that we observed were on the ground within a few centimeters of the water. The males usually are hidden beneath an overhanging leaf or some other cover; they definitely do not sit in the open likeSmilisca baudini. Most calling males and clasping pairs have been found at the edges of small pools or shallow ditches, although occasional individuals are found at the edges of large ponds or streams.The breeding call consists of one or two moderately short, low-pitched notes (duration 0.33 to 0.42 seconds), repeated at intervals of about 20 seconds to several minutes. Each note is a low, vibrant "wauk," having 100 to 130 pulses per second and a dominant frequency of 330 to 420 cycles per second (Pl. 10C).The eggs are deposited in loose clumps amidst vegetation in the water. Hatchling tadpoles have total lengths of 8.7 to 10.6 mm., and body lengths of 4.1 to 4.5 mm. The external gills are long and filamentous, and the yolk sac is large. The head and caudal musculature are dark brownish black, and the caudal fins are gray. The oral discs are large and roughly circular. The growth and development of the tadpoles are summarized in table 11 and figure 16.A typical tadpole in stage 30 of development (KU 68482 from the Río Chitaría, Cartago Province, Costa Rica) may be described as follows: body length 9.7 mm.; tail length 14.6 mm.; total length 24.3 mm.; body as wide as deep; snout rounded dorsally and laterally; eyes widely separated, directeddorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, about midway on length of body and slightly below midline; anal tube dextral; caudal musculature slender, curved upward distally; dorsal fin extending onto body; depth of dorsal fin slightly less than that of ventral fin at mid-length of tail; dorsal part of body pale brown; ventral surfaces transparent with scattered pigment; pale cream-colored, crescent-shaped mark on posterior edge of body; caudal musculature pale creamy tan with scattered pale brown spots; caudal fins transparent with scattered small brown blotches on dorsal and ventral fins; iris pale bronze in life (Fig. 13); mouth small; median part of upper lip bare; rest of mouth bordered by one row of pointed papillae; lateral fold present; tooth-rows 2/3, first upper row longest; second upper row slightly shorter, broadly interrupted medially; three lower rows complete, equal in length, slightly shorter than second upper row; upper beak moderately deep, forming broad arch with slender lateral processes; lower beak slender, broadly V-shaped; both beaks serrate (Fig. 15E).In tadpoles having fully developed mouthparts the tooth-row formula of 2/3 is invariable. The pale crescent-shaped mark on the posterior part of the body curves anterodorsally on the dorsal surface of the body. These marks in dorsal view give the appearance of a pair of short, pale stripes on the posterior part of the body. Most specimens from Costa Rica have the pale coloration like that described above, but some individuals (notable KU 87683 from Guápiles, Costa Rica, KU 87707 from Finca Tepeyac, Nicaragua, and KU 87708 from Bonanza, Nicaragua) have much more pigment. In these specimens the same color pattern obtains as in the pallid individuals, but the pigmentation is dense. This is especially noticeable on the tail.Recently metamorphosed young have snout-vent lengths of 12.7 to 16.7 mm. (average, 14.3 mm. in eleven specimens). Coloration of young in life (KU 68484 from Río Chitaría, Cartago Province, Costa Rica): "Dorsum pale tan; side of head and flanks darker brown, separated from tan dorsum by an indistinct cream stripe. Limbs pale yellow; thighs flecked with brown; shank and tarsus yellowish tan with indistinct brown bars. Soles of feet brown. Belly white; throat dusty cream flecked with silvery white. Upper lip silvery white. Iris bright gold with black flecks. Heels, tarsal and anal stripes white" (Duellman, field notes, May 23, 1961).Remarks.—Peters (1863:463) namedHyla labialisfrom the "region of Bogotá, Colombia", but in 1873 regarded his new species as identical withHyla phaeotaCope, 1862, from Turbo, Colombia. The nameHyla labialishas been used for frogs from the northern Andes in Colombia (see Dunn, 1944:72, and Stebbins and Hendrickson, 1959:522, for discussion of nomenclature). Rivero (1961:131) used the nameHyla vilsonianaCope, 1899, for the frogs from the northern Andes previously referred toHyla labialis. A review of the nomenclature and taxonomy of these frogs, which superficially resembleSmiliscabut are unrelated, is beyond the scope of the present study.Hyla baudini dolomedesBarbour, 1923, is based on a smallSmilisca phaeota(MCZ 8539) having a snout-vent length of 45.5 mm. Dunn (1931a:413) placeddolomedesin the synonymy ofSmilisca phaeota. We have examined the holotype ofdolomedesand agree with Dunn's assignment.Smith (1953:150) describedHyla phaeota cyanostictafrom Guatemala. Our studies on the external morphology, coloration, and especially the cranial osteology provide evidence thatcyanostictais a species distinct fromphaeota.Distribution.—Smilisca phaeotainhabits 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 Panamá, 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 Río Cauca and Río 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; Río Mico, 16 km. E Recreo, UMMZ 79711 (6), 79712 (4); junction Río Mico and Río Siguia, UMMZ 79713 (10); Río 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; Río Chitaría, 3 km. NNE Pavones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); Río Reventazón, 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: Tilarán, KU 36805-7; 8 km. NE Tilarán, KU 36803-4.Heredia: Barranca del Río Sarapiquí 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.Limón: Bambú, USC 7182 (4); Batán, UMMZ 118582; Coén, 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 Limón, KU 32261; Río Larí at Río Dipari, 21 km. SW Amubre, USC 7177; Río Toro Amarillo, 7 km. W Guápiles, 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; Rincón, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), USC 7254; Río Ferruviosa, 7 km. S Rincón, USC 7235; 1.6 km. WNW Villa Neily, KU 68493 (young), 68494 (tadpoles).San José: 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.Coclé: El Valle, KU 77521-4, 77649 (tadpole), TNHC 23369.Comarca del Barú: Progreso, UMMZ 61085-9. Colón: Achiote, KU 77516-20, 77648 (young); Río Candelaria, CNHM 67851-2.Darién: Río Esnápe, Sambú Valley, MCZ 8539; Río Sucubti, Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, Camp Townsend, AMNH 40988.Panamá: NW slope Cerro Prominente, KU 80459; Finca La Sumbadora, KU 91914 (skeleton).Chiriquí: 2 km. W Concepción, 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 Urabá, CNHM 63881 (Goin); Peña Lisa, Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro, CNHM 4451-3, 4455-61 (Goin); Río San Juan, Playa del Oro, CNHM 54772 (Goin); Río Quesada, AMNH 13615-77; 37 km. up Río Puné, AMNH 13688; 48 km. up Río Puné, AMNH 13689.Narino: Tumaco, Río Rosario, CJG 2310-13 (Goin).Valle: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT 166, 346-47, 388 (Goin); Río Calima above Córdoba, CJG 2249-57 (Goin).Ecuador: No province: Bulun, AMNH 10620.Esmeraldas: Cachabé, AMNH 10625-8; Río Capayas, CNHM 35712; Río 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 combinationHyla pumaCope, Proc. Amer. Philos. Soc., 22:183, 1885 [Holotype.—USNM 13735 from Nicaragua; J. F. Moser collector]. Günther, 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 wellmanorumTaylor, Univ. Kansas Sci. Bull. 25(1):843, July 1, 1952 [Holotype.—KU 30302 from Batán, Limón, 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 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 distinguishingS. pumafrom 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 pumainhabits 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 Batán, 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, 0.06-0.35seconds; 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'sHyla pumahas 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 ofSmilisca wellmanorum(Taylor, 1952); the vestigial webbing on thehands and the dorsal coloration are especially significant. Consequently, we considerHyla wellmanorumTaylor, 1952, to be a synonym ofHyla pumaCope, 1885. Cope gave only "Nicaragua" as the locality forHyla 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 areDentrobates 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 ofHyla pumacame 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, listedHyla pumaCope, 1885, as a synonym ofHyla molitorSchmidt, 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 calledMueller'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 colorationHyla pumais distinctly different fromHyla 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.Fig. 3.Map showing locality records forSmilisca puma(triangles) andSmilisca 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); Río 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 PuertoViejo, 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: Batán, 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 silanew speciesHyla 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 Volcán, Chiriquí Province, Panamá, 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 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 distinguishingS. silafrom 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[Pg 319]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, Panamá, 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 Panamá). 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 Volcán, Chiriquí, Panamá, 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 Volcán 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 Panamá. The truncate snout is characteristic of both sexes.The coloration ofSmilisca silaconsists 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 inSmilisca sordida. In one specimen (KU 80467) the blotches are fused and form two wide irregular longitudinal stripes, as inSmilisca 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 Panamá have distinct bars on the limbs; the posterior surfaces of the thighs have brown reticulations enclosing small blue flecks in specimens fromCosta Rica and bolder, black reticulations enclosing large pale blue spots in specimens from western Panamá. In specimens from Costa Rica the flanks are brown with pale blue flecks, whereas in those from Chiriquí, Panamá, 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 Chiriquí, 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 Darién and from Panamá Province east of the Canal Zone (Altos de Pacora, Cerro Jefe, Finca La Sumbadora, and Río 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, Panamá, 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, Panamá, 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 Panamá west of the Canal Zone, the blue coloration on the flanks and thighs is much less conspicuous than in specimens from eastern Panamá. 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, Chiriquí, Panamá, 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 silainhabits 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 Volcán, Chiriquí, Panamá). 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, Panamá) 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; musculatureextending 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 Volcán, Chiriquí (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.Metamorphosing young have been found on vegetation at the edge of streams and have been raised in the laboratory. Seven recently metamorphosed young have snout-vent lengths of 13.6 to 15.6 mm. (14.6 mm.). A living juvenile (KU 91913) raised in the laboratory from a tadpole obtained at Finca La Sumbadora had a brown dorsum with darker brown markings, a white spot below the eye, and a narrow white labial stripe. The belly was white; the flanks were brown with white spots, and the posterior surfaces of the thighs were yellowish tan. The iris was a golden bronze color with much black reticulation.Remarks.—This species has been confused withSmilisca sordida; most authors have referred both species toHyla (Smilisca) gabbi. Examination of the types ofHyla sordida,gabbi,salvini, andnigripesrevealed that all of the names were referable to a single species (S. sordida), and that the small, blunt-snouted species in Panamá and southern Costa Rica probably was without a name. PossiblyHyla molitorSchmidt (1857) is based on the species that we have namedS. sila, but several discrepancies in his description, plus the unknown provenance of the type, have led us to discount the applicability of that name to the species under consideration.Distribution.—Smilisca silaranges along the Pacific slopes and lowlands of southern Costa Rica and Panamá at elevations from sea level to about 1300 meters; in northern South America the species occurs in theCaribbeanlowlands and in the valleys of the northward draining rivers of Colombia (Fig. 3).
Specimens examined.—78, as follows:Mexico:Chiapas: Monte Libano, MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH 41588.Oaxaca: 11 km. N Vista Hermosa, KU 84918-20 (skeletons), 87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH 57199-201; 8 km. S Yetla, KU 87213, UMMZ 124838 (8).Veracruz: Coyame, UMMZ 111459-60; between Coyame and Tebanco, UMMZ 121198; Dos Amates, UMMZ 121297; between Laguna de Catemaco and Volcán San Martín, UMMZ 121200; Volcán San Martín, UIMNH 35403-4, 35408-12, UMMZ 118163; SE slope Volcán San Martín, UMMZ 121199, 121295 (2), 121296, 121298.Guatemala:Alta Verapaz: Chinajá, KU 55935-7, 55938 (skeleton).El
Specimens examined.—78, as follows:Mexico:Chiapas: Monte Libano, MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH 41588.Oaxaca: 11 km. N Vista Hermosa, KU 84918-20 (skeletons), 87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH 57199-201; 8 km. S Yetla, KU 87213, UMMZ 124838 (8).Veracruz: Coyame, UMMZ 111459-60; between Coyame and Tebanco, UMMZ 121198; Dos Amates, UMMZ 121297; between Laguna de Catemaco and Volcán San Martín, UMMZ 121200; Volcán San Martín, UIMNH 35403-4, 35408-12, UMMZ 118163; SE slope Volcán San Martín, UMMZ 121199, 121295 (2), 121296, 121298.
Guatemala:Alta Verapaz: Chinajá, KU 55935-7, 55938 (skeleton).El
Petén: 10 km. NNW Chinajá (Alta Verapaz), KU 55934; Piedras Negras, CNHM 99006-7, 99227, UIMNH 28853, USNM 111139-41, 111143-7; 8 km. S Piedras Negras, CNHM 99008; Semicoch, USNM 35907.
Fig. 2.Map showing locality records forSmilisca cyanosticta(triangles) andSmilisca phaeota(circles).
Fig. 2.Map showing locality records forSmilisca cyanosticta(triangles) andSmilisca phaeota(circles).
Smilisca phaeota(Cope)
Hyla phaeotaCope, Proc. Acad. Nat. Sci. Philadelphia, 14 (9):358, 1862 [Holotype.—USNM 4347 from Turbo, Colombia; J. Cassin collector]. Boulenger, Catalogue Batrachia Salientia in British Museum, p. 402, Feb. 1, 1882. Werner, Sitzungsb. Akad. Wiss. München, 27:215, 1897. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p. 269, Sept. 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 261, June 1923. Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, Oct. 10, 1931. Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 357:4, Oct. 26, 1937. Cooper, Copeia, 2:122, June 30, 1944. Breder, Bull. Amer. Mus. Nat. Hist., 86(6):416, Aug. 26, 1946. Smith and Taylor, Bull. U.S. Natl. Mus., 194:88, June 17, 1948; Univ. Kansas Sci. Bull, 33:364, March 20, 1950. Taylor, Univ. Kansas Sci. Bull., 35(1):837, July 1, 1952. Brattstrom and Howell, Herpetologica, 10:117,[Pg 309]Aug. 1, 1954. Goin, Herpetologica, 14:120, July 23, 1958. Cochran, Bull. U.S. Natl. Mus., 220:57, 1961.
Hyla labialisPeters, Monats. Konigl. Akad. Wissen. Berlin, p. 463, 1863 [Holotype.—ZMB 4913 from "region of Bogotá," Colombia]; Monats. Konigl. Akad. Wissen. Berlin, p. 618, Oct. 16, 1873. Boulenger, Catalogue Batrachia and Salientia in British Museum, p. 397, Feb. 1, 1882.
Hyla baudini dolomedesBarbour, Occas. Papers Mus. Zool. Univ. Michigan, 129:11, Jan. 25, 1923 [Holotype.—MCZ 8539 from Río Esnápe, Sambú Valley, Darién, Panamá; Barbour and Brooks collectors]. Barbour and Loveridge, Bull. Mus. Comp. Zool. Harvard, 69:278, June, 1929.
Hyla phaeota phaeota, Smith, Herpetologica, 8:152, Jan. 30, 1953. Minton and Smith, Herpetologica, 16:103, June 17, 1960.
Smilisca phaeota, Starrett, Copeia, 4:303, Dec. 30, 1960.
Diagnosis.—Size large ([M] 65 mm., [F] 78 mm.); skull as long as wide, lacking frontoparietal fontanelle; large supraorbital flanges having straight edges and extending posterolaterally; large squamosal not in contact with maxillary; tarsal fold moderately wide, full length of tarsus; inner metatarsal tubercle moderately large, low, flat, elliptical; hind limbs relatively long, tibia averaging more than 53 per cent of snout-vent length; labial stripe silvery white; lips lacking vertical bars; loreal region pale green; dark brown or black tympanic mark dispersing into brown venated pattern on flanks; posterior surfaces of thighs pale brown, with or without darker flecks or small cream-colored flecks. (Foregoing combination of characters distinguishingS. phaeotafrom any other species in genus.)
Description and Variation.—For the purposes of analyzing geographic variation in size and proportions, measurements were taken on ten adult males from each of five samples throughout the range of the species. Aside from the data summarized in Table 2, the average ratio of tibia length to snout-vent length is noticeably less in Colombian specimens (53.4 per cent, as compared with 54.8 to 57.8 per cent in the other samples) and the ratio of head length tosnout-vent length is noticeably less in Costa Rican specimens (33.5 per cent as compared with 34.9 to 35.1 per cent in the other samples). Also, specimens from Heredia Province, Costa Rica, have a relatively smaller tympanum (62.7 to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared with means of 74.0 to 77.9 per cent in the other samples).
Two populations are distinctive as regards the size of adult males. Specimens from the northern Caribbean lowlands of Nicaragua (Bonanza, the northernmost locality for the species) are remarkably small. Males having snout-vent lengths of between 40 and 43 mm. were breeding; the largest male found had a snout-vent length of 47.7 mm. The other extreme in size is attained in specimens from the Pacific lowlands of eastern Costa Rica and western Panamá, where most breeding males have snout-vent lengths of more than 55 mm.; the largest male had a snout-vent length of 64.9 mm.
The rather striking differences in size among certain samples and the minor differences in proportions among other samples show no geographic trends. Instead, the variations apparently are random among the samples. The data presented here possibly are the results of inadequate sampling, but more likely reflect actual differences in the populations.
The dorsal ground color ofSmilisca phaeotais pale green to tan; the venter is creamy white. The dorsum is variously marked with dark olive-green or dark brown spots or blotches (Pl. 6C). A dark interorbital bar is usually present. Usually a large dark dorsal mark extends from the occiput to the sacral region, but in many individuals this blotch is replaced by two or three dark marks. The dorsal markings are irregular in shape and do not tend to form transverse bands or longitudinal bars. The hind limbs are marked by dark transverse bands, usually four or five on the thigh, five or six on the shank, and four on the tarsus. Two or three narrow bands are usually present on the proximal part of the fourth toe. The webbing on the feet is brown. The loreal region is pale green, bordered above by a narrow dark brown canthal stripe extending from the nostril to the orbit. The upper lip is silvery white. A broad dark brown or black mark extends posteriorly from the orbit, encompassing the tympanum, to a point above the insertion of the forelimb. The flanks are pale green or pale tan and marked with a fine dark brown or black venation. The anterior surfaces of the thighs usually are pale brown or grayish tan, sometimes having small, indistinct darker flecks. The posterior surfaces of the thighs are similarly colored, but in most specimens small but distinct dark flecks are present; in some specimens small cream-colored spots are also present on the posterior surfaces of the thighs. A distinct, narrow creamy white anal stripe usually is present. A distinct white stripe is present on the outer edge of the tarsus and fifth toe; on the tarsus the white stripe is bordered below by dark brown. A white stripe also is present on the outer edge of the forearm and fourth finger. In breeding males the throat is dark gray.
Little geographic variation in color or pattern is evident. Few, if any, specimens from the Pacific lowlands of South America are green in life. (We have seen no living individuals from South America.) Some living individuals from Costa Rica and all those seen alive from Nicaragua have a tint of pale blue on the flanks. In some specimens the dorsal pattern is so faint as to be barely discernible, whereas in most specimens the pattern is bold.
The coloration in the living frogs is highly variable due to extreme metachrosis. Individuals of this species are capable of changing the dorsal colorationfrom green to brown in a short period of time. Both green and brown individuals have been found active at night. Usually those individuals found hiding by day are brown. One individual from Finca La Sumbadora, Panamá (now KU 91914), was kept alive in the laboratory for nearly one month. This individual usually was pale green with tan dorsal markings at night and tan with pale green markings by day. On occasion the pale green dorsal markings were boldly outlined by bright dark green.
In living individuals from throughout the range of the species the iris is a bronze color, darkest medially with fine black reticulations.
Natural History.—Smilisca phaeotainhabits humid lowland tropical forest and seldom ascends the foothills to more than 1,000 meters. The rather equable climatic conditions, especially more or less evenly distributed rainfall throughout the year, permit this frog to be active most of the year. Dunn (1931:413) reported males calling on Barro Colorado Island, Panamá, in February and in July, and Breder (1946:416) noted calling individuals in the Chucanaque drainage of Darién, Panamá in January, March, July, August and October and in Costa Rica in April through August inclusively. Calling males were found at Bonanza, Nicaragua in March and in July.
At all times of year the usual daytime retreats for these frogs are near water; the frogs have been found in elephant ear plants (Xanthosoma) and in bromeliads; occasional individuals have been found sitting on shaded branches of bushes and trees. None has been observed on the ground or beneath ground-cover by day.
The length of the breeding season cannot be determined definitely. The earliest date on which eggs have been found is May 23; Gaige, Hartweg, and Stuart (1937:5) reported a gravid female taken at El Recreo, Nicaragua, in September, and we have a gravid female taken at Almirante, Panamá, in March.
Males usually call from secluded spots at the edge of water. All calling males that we observed were on the ground within a few centimeters of the water. The males usually are hidden beneath an overhanging leaf or some other cover; they definitely do not sit in the open likeSmilisca baudini. Most calling males and clasping pairs have been found at the edges of small pools or shallow ditches, although occasional individuals are found at the edges of large ponds or streams.
The breeding call consists of one or two moderately short, low-pitched notes (duration 0.33 to 0.42 seconds), repeated at intervals of about 20 seconds to several minutes. Each note is a low, vibrant "wauk," having 100 to 130 pulses per second and a dominant frequency of 330 to 420 cycles per second (Pl. 10C).
The eggs are deposited in loose clumps amidst vegetation in the water. Hatchling tadpoles have total lengths of 8.7 to 10.6 mm., and body lengths of 4.1 to 4.5 mm. The external gills are long and filamentous, and the yolk sac is large. The head and caudal musculature are dark brownish black, and the caudal fins are gray. The oral discs are large and roughly circular. The growth and development of the tadpoles are summarized in table 11 and figure 16.
A typical tadpole in stage 30 of development (KU 68482 from the Río Chitaría, Cartago Province, Costa Rica) may be described as follows: body length 9.7 mm.; tail length 14.6 mm.; total length 24.3 mm.; body as wide as deep; snout rounded dorsally and laterally; eyes widely separated, directeddorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, about midway on length of body and slightly below midline; anal tube dextral; caudal musculature slender, curved upward distally; dorsal fin extending onto body; depth of dorsal fin slightly less than that of ventral fin at mid-length of tail; dorsal part of body pale brown; ventral surfaces transparent with scattered pigment; pale cream-colored, crescent-shaped mark on posterior edge of body; caudal musculature pale creamy tan with scattered pale brown spots; caudal fins transparent with scattered small brown blotches on dorsal and ventral fins; iris pale bronze in life (Fig. 13); mouth small; median part of upper lip bare; rest of mouth bordered by one row of pointed papillae; lateral fold present; tooth-rows 2/3, first upper row longest; second upper row slightly shorter, broadly interrupted medially; three lower rows complete, equal in length, slightly shorter than second upper row; upper beak moderately deep, forming broad arch with slender lateral processes; lower beak slender, broadly V-shaped; both beaks serrate (Fig. 15E).
In tadpoles having fully developed mouthparts the tooth-row formula of 2/3 is invariable. The pale crescent-shaped mark on the posterior part of the body curves anterodorsally on the dorsal surface of the body. These marks in dorsal view give the appearance of a pair of short, pale stripes on the posterior part of the body. Most specimens from Costa Rica have the pale coloration like that described above, but some individuals (notable KU 87683 from Guápiles, Costa Rica, KU 87707 from Finca Tepeyac, Nicaragua, and KU 87708 from Bonanza, Nicaragua) have much more pigment. In these specimens the same color pattern obtains as in the pallid individuals, but the pigmentation is dense. This is especially noticeable on the tail.
Recently metamorphosed young have snout-vent lengths of 12.7 to 16.7 mm. (average, 14.3 mm. in eleven specimens). Coloration of young in life (KU 68484 from Río Chitaría, Cartago Province, Costa Rica): "Dorsum pale tan; side of head and flanks darker brown, separated from tan dorsum by an indistinct cream stripe. Limbs pale yellow; thighs flecked with brown; shank and tarsus yellowish tan with indistinct brown bars. Soles of feet brown. Belly white; throat dusty cream flecked with silvery white. Upper lip silvery white. Iris bright gold with black flecks. Heels, tarsal and anal stripes white" (Duellman, field notes, May 23, 1961).
Remarks.—Peters (1863:463) namedHyla labialisfrom the "region of Bogotá, Colombia", but in 1873 regarded his new species as identical withHyla phaeotaCope, 1862, from Turbo, Colombia. The nameHyla labialishas been used for frogs from the northern Andes in Colombia (see Dunn, 1944:72, and Stebbins and Hendrickson, 1959:522, for discussion of nomenclature). Rivero (1961:131) used the nameHyla vilsonianaCope, 1899, for the frogs from the northern Andes previously referred toHyla labialis. A review of the nomenclature and taxonomy of these frogs, which superficially resembleSmiliscabut are unrelated, is beyond the scope of the present study.
Hyla baudini dolomedesBarbour, 1923, is based on a smallSmilisca phaeota(MCZ 8539) having a snout-vent length of 45.5 mm. Dunn (1931a:413) placeddolomedesin the synonymy ofSmilisca phaeota. We have examined the holotype ofdolomedesand agree with Dunn's assignment.
Smith (1953:150) describedHyla phaeota cyanostictafrom Guatemala. Our studies on the external morphology, coloration, and especially the cranial osteology provide evidence thatcyanostictais a species distinct fromphaeota.
Distribution.—Smilisca phaeotainhabits 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 Panamá, 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 Río Cauca and Río 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; Río Mico, 16 km. E Recreo, UMMZ 79711 (6), 79712 (4); junction Río Mico and Río Siguia, UMMZ 79713 (10); Río 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; Río Chitaría, 3 km. NNE Pavones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); Río Reventazón, 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: Tilarán, KU 36805-7; 8 km. NE Tilarán, KU 36803-4.Heredia: Barranca del Río Sarapiquí 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.Limón: Bambú, USC 7182 (4); Batán, UMMZ 118582; Coén, 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 Limón, KU 32261; Río Larí at Río Dipari, 21 km. SW Amubre, USC 7177; Río Toro Amarillo, 7 km. W Guápiles, 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; Rincón, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), USC 7254; Río Ferruviosa, 7 km. S Rincón, USC 7235; 1.6 km. WNW Villa Neily, KU 68493 (young), 68494 (tadpoles).San José: 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.Coclé: El Valle, KU 77521-4, 77649 (tadpole), TNHC 23369.Comarca del Barú: Progreso, UMMZ 61085-9. Colón: Achiote, KU 77516-20, 77648 (young); Río Candelaria, CNHM 67851-2.Darién: Río Esnápe, Sambú Valley, MCZ 8539; Río Sucubti, Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, Camp Townsend, AMNH 40988.Panamá: NW slope Cerro Prominente, KU 80459; Finca La Sumbadora, KU 91914 (skeleton).Chiriquí: 2 km. W Concepción, 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 Urabá, CNHM 63881 (Goin); Peña Lisa, Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro, CNHM 4451-3, 4455-61 (Goin); Río San Juan, Playa del Oro, CNHM 54772 (Goin); Río Quesada, AMNH 13615-77; 37 km. up Río Puné, AMNH 13688; 48 km. up Río Puné, AMNH 13689.Narino: Tumaco, Río Rosario, CJG 2310-13 (Goin).Valle: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT 166, 346-47, 388 (Goin); Río Calima above Córdoba, CJG 2249-57 (Goin).Ecuador: No province: Bulun, AMNH 10620.Esmeraldas: Cachabé, AMNH 10625-8; Río Capayas, CNHM 35712; Río 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.
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; Río Mico, 16 km. E Recreo, UMMZ 79711 (6), 79712 (4); junction Río Mico and Río Siguia, UMMZ 79713 (10); Río 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; Río Chitaría, 3 km. NNE Pavones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); Río Reventazón, 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: Tilarán, KU 36805-7; 8 km. NE Tilarán, KU 36803-4.
Heredia: Barranca del Río Sarapiquí 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.
Limón: Bambú, USC 7182 (4); Batán, UMMZ 118582; Coén, 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 Limón, KU 32261; Río Larí at Río Dipari, 21 km. SW Amubre, USC 7177; Río Toro Amarillo, 7 km. W Guápiles, 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; Rincón, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), USC 7254; Río Ferruviosa, 7 km. S Rincón, USC 7235; 1.6 km. WNW Villa Neily, KU 68493 (young), 68494 (tadpoles).
San José: 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.Coclé: El Valle, KU 77521-4, 77649 (tadpole), TNHC 23369.Comarca del Barú: Progreso, UMMZ 61085-9. Colón: Achiote, KU 77516-20, 77648 (young); Río Candelaria, CNHM 67851-2.Darién: Río Esnápe, Sambú Valley, MCZ 8539; Río Sucubti, Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, Camp Townsend, AMNH 40988.Panamá: NW slope Cerro Prominente, KU 80459; Finca La Sumbadora, KU 91914 (skeleton).Chiriquí: 2 km. W Concepción, 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 Urabá, CNHM 63881 (Goin); Peña Lisa, Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro, CNHM 4451-3, 4455-61 (Goin); Río San Juan, Playa del Oro, CNHM 54772 (Goin); Río Quesada, AMNH 13615-77; 37 km. up Río Puné, AMNH 13688; 48 km. up Río Puné, AMNH 13689.Narino: Tumaco, Río Rosario, CJG 2310-13 (Goin).Valle: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT 166, 346-47, 388 (Goin); Río Calima above Córdoba, CJG 2249-57 (Goin).
Ecuador: No province: Bulun, AMNH 10620.Esmeraldas: Cachabé, AMNH 10625-8; Río Capayas, CNHM 35712; Río 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 pumaCope, Proc. Amer. Philos. Soc., 22:183, 1885 [Holotype.—USNM 13735 from Nicaragua; J. F. Moser collector]. Günther, 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 wellmanorumTaylor, Univ. Kansas Sci. Bull. 25(1):843, July 1, 1952 [Holotype.—KU 30302 from Batán, Limón, 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 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 distinguishingS. pumafrom 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 pumainhabits 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 Batán, 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, 0.06-0.35seconds; 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'sHyla pumahas 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 ofSmilisca wellmanorum(Taylor, 1952); the vestigial webbing on thehands and the dorsal coloration are especially significant. Consequently, we considerHyla wellmanorumTaylor, 1952, to be a synonym ofHyla pumaCope, 1885. Cope gave only "Nicaragua" as the locality forHyla 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 areDentrobates 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 ofHyla pumacame 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, listedHyla pumaCope, 1885, as a synonym ofHyla molitorSchmidt, 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 calledMueller'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 colorationHyla pumais distinctly different fromHyla 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.
Fig. 3.Map showing locality records forSmilisca puma(triangles) andSmilisca sila(circles).
Fig. 3.Map showing locality records forSmilisca puma(triangles) andSmilisca 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); Río 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 PuertoViejo, 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: Batán, 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).
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); Río 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 PuertoViejo, 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: Batán, 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 silanew 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 Volcán, Chiriquí Province, Panamá, 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 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 distinguishingS. silafrom 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[Pg 319]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 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[Pg 319]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, Panamá, 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 Panamá). 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 Volcán, Chiriquí, Panamá, 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 Volcán 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 Panamá. The truncate snout is characteristic of both sexes.
The coloration ofSmilisca silaconsists 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 inSmilisca sordida. In one specimen (KU 80467) the blotches are fused and form two wide irregular longitudinal stripes, as inSmilisca 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 Panamá have distinct bars on the limbs; the posterior surfaces of the thighs have brown reticulations enclosing small blue flecks in specimens fromCosta Rica and bolder, black reticulations enclosing large pale blue spots in specimens from western Panamá. In specimens from Costa Rica the flanks are brown with pale blue flecks, whereas in those from Chiriquí, Panamá, 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 Chiriquí, 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 Darién and from Panamá Province east of the Canal Zone (Altos de Pacora, Cerro Jefe, Finca La Sumbadora, and Río 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, Panamá, 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, Panamá, 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 Panamá west of the Canal Zone, the blue coloration on the flanks and thighs is much less conspicuous than in specimens from eastern Panamá. 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, Chiriquí, Panamá, 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 silainhabits 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 Volcán, Chiriquí, Panamá). 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, Panamá) 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; musculatureextending 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 Volcán, Chiriquí (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.
Metamorphosing young have been found on vegetation at the edge of streams and have been raised in the laboratory. Seven recently metamorphosed young have snout-vent lengths of 13.6 to 15.6 mm. (14.6 mm.). A living juvenile (KU 91913) raised in the laboratory from a tadpole obtained at Finca La Sumbadora had a brown dorsum with darker brown markings, a white spot below the eye, and a narrow white labial stripe. The belly was white; the flanks were brown with white spots, and the posterior surfaces of the thighs were yellowish tan. The iris was a golden bronze color with much black reticulation.
Remarks.—This species has been confused withSmilisca sordida; most authors have referred both species toHyla (Smilisca) gabbi. Examination of the types ofHyla sordida,gabbi,salvini, andnigripesrevealed that all of the names were referable to a single species (S. sordida), and that the small, blunt-snouted species in Panamá and southern Costa Rica probably was without a name. PossiblyHyla molitorSchmidt (1857) is based on the species that we have namedS. sila, but several discrepancies in his description, plus the unknown provenance of the type, have led us to discount the applicability of that name to the species under consideration.
Distribution.—Smilisca silaranges along the Pacific slopes and lowlands of southern Costa Rica and Panamá at elevations from sea level to about 1300 meters; in northern South America the species occurs in theCaribbeanlowlands and in the valleys of the northward draining rivers of Colombia (Fig. 3).