Chapter II: Part 2

=Tamaulipas=: Acuna, UMMZ 99864; 5 km. S Acuna, UMMZ 101180; 13 km. N Antiguo Morelos, UIMNH 40532-5; 3 km. S Antiguo Morelos, UF 11204; 3 km. NE Chamal, UMMZ 102867; 1.6 km. E Chamal, UMMZ 110734; Ciudad Mante, UMMZ 80957, 80958 (3), 106400 (3); 16 km. N Ciudad Victoria, CNHM 102408; 34 km. NE Ciudad Victoria, KU 60395-411; 8.8 km. S Ciudad Victoria, UIMNH 19261-3; 11 km. W Ciudad Victoria, UIMNH 30924; 16 km. W Ciudad Victoria, UIMNH 30825; 3 km. W El Carizo, UMMZ 111279; Gomez Farias, UMMZ 110837-8; 8 km. NE Gomez Farias, UMMZ 102265, 102916 (4), 102917, 104110 (5), 105493, 110836 (2), 111274-7; 8 km. NW Gomez Farias, UMMZ 101178 (7), 101179 (3), 101362-3, 101364 (2), 108799 (2), 110129, 111278, 111280; 8 km. W Gomez Farias, UMMZ 102859 (2); 16 km. W Gonzales, KU 37795-6; Jimenez, KU 60412; La Clementina, 6 km. W Forlan, USNM 106244; Limon, UIMNH 30831; Llera, USNM 140137-40; 3 km. E Llera, UIMNH 16858; 21 km. S Llera, UIMNH 30828-9; 23 km. S Llera, UIMNH 30830; 11 km. SW Ocampo, UMMZ 118956; 22 km. W, 5 km. S Piedra, KU 37568-71; Rio Sabinas, UMMZ 97976; 5 km. W San Gerardo, UMMZ 110733 (2); Santa Barbara, UMMZ 111272-3; Villagran, CNHM 102280, 102287, 102299, 102309, UIMNH 30826-7; 1.7 km. W Xicotencatl, UMMZ 115179.

=Veracruz=: 1.6 km. NW Acayucan, UMMZ 115189; 28.5 km. SE Alvarado, UMMZ 119933; 2.4 km. SSW Amatitlan, UMMZ 115195; Barranca Metlac, UIMNH 38365; Boca del Rio, UIMNH 26619-30, UMMZ 74954 (9); 16 km. S Boca del Rio, UIMNH 26631; between Boca del Rio and Anton Lizardo, UIMNH 42701; Canada, CNHM 102397; Catemaco, UMMZ 118702 (4); Ciudad Aleman, UMMZ 119608 (3); Cordoba, CNHM 38665-7, USNM 30410-3; 5.2 km. ESE Cordoba, KU 71423-35, 89924 (skeleton); 7 km. ESE Cordoba, UMMZ 115176 (4); Cosamaloapan, UMMZ 115193 (2); Coyame, UIMNH 36853-6, 38366, UMMZ 111461 (3), 111462-3; 1 km. SE Coyame, UMMZ 121202 (3); Cuatotolapam, UMMZ 41625-39; Cuautlapan, CNHM 38664, 70591-600, 102218, 102398, KU 26300, 26302, 26309, 26312-3, 26315-6, 26321, 26336, 26339, 26347 (skeleton), 26354, 55614-21 (skeletons), UIMNH 11236-67, 11269-71, 11273, 26611-8, 30792-5, UMMZ 85466 (6), 115173 (25), 115175 (7), USNM 114433-57; Dos Rios, CNHM 39623; 5 km. ENE El Jobo, KU 23843, 23845, 23847; 6.2 km. E Encero, UIMNH 30835; Escamilo, CNHM 102298, UIMNH 30788; 1 km. N Fortin, UF 11201; 4 km. SW Huatusco, UMMZ 115177; 1 km. SW Huatusco, UMMZ 123119; 10 km. SE Hueyapan, UMMZ 115190; 20 km. S Jesus Carranza, KU 23844, 23846, 27399; 38 km. SE Jesus Carranza, KU 23417; Laguna Catemaco, UMMZ 119932 (62); 1.6 km. N La Laja, UIMNH 3651; La Oaxaquena, AMNH 43930-1; 17 km. E Martinez de la Torre, UIMNH 36630-2; 6.2 km. W Martinez de la Torre, UIMNH 3652-4; Minatitlan, AMNH 52141-2; Mirador, USNM 25097-8, 115178; 6 km. S Monte Blanco, UF 11200 (4); 21 km. E Nanchital, UMMZ 123004; 2 km. S Naranja, UMMZ 115188 (3); 1.6 km. NE Novillero, UMMZ 115194 (2); 3 km. NE Novillero, UMMZ 115196; 5.2 km. NE Novillero, UMMZ 115192 (4); 6 km. NE Novillero, UMMZ 115191; 5 km. N Nuevo Colonia, UMMZ 105066; Orizaba, USNM 16563-6; 4 km. NE Orizaba, UMMZ 120251 (2); Panuco, UMMZ 118922; Paraje Nuevo, UMMZ 85465 (2), 85467 (35), 85468 (36); Paso del Macho, UIMNH 49281; Paso de Talaya, Jicaltepec, USNM 32365, 84420; Perez, CNHM 1686 (5); 20 km. N Piedras Negras, Rio Blanco, KU 23708; Plan del Rio, KU 26310, 26333-5, 26345, 26354, UMMZ 102069, 102070 (5); Potrero, UIMNH 49282-5, UMMZ 88799, 88805, 88806 (2), USNM 32391-5; Potrero Viejo, CNHM 102296, KU 26301, 26304-5, 26307-8, 26311, 26317-20, 26323-25, 26326-8 (skeletons), 26329-31, 26332 (skeleton), 26337-8, 26340-4, 26346, 26348, 26351, 26353, 27400-12, UIMNH 30800, UMMZ 88800 (2), 88802 (15), 88803 (9), 88804, USNM 114458-67; 5 km. S Potrero Viejo, KU 26303, 26314, 26322; Puente Nacional, UIMNH 21783-8; 3 km. N Rinconada, UMMZ 122099 (5); Rio de las Playas, USNM 118635-6; Rio Seco, UMMZ 88801 (9); Rodriguez Clara, CNHM 102225; San Andres Tuxtla, CNHM 102213, 102222, 102227, 102247, UIMNH 30789-91; 10 km. NW San Andres Tuxtla, UMMZ 119935; 13.4 km. NW San Andres Tuxtla, UMMZ 119939 (2); 19.8 km. NW San Andres Tuxtla, UMMZ 119938; 27.2 km. NW San Andres Tuxtla, UMMZ 119936 (6); 48 km. NW San Andres Tuxtla, UMMZ 119937; 4 km. W San Andres Tuxtla, UMMZ 115187; 37.4 km. S San Andres Tuxtla, UMMZ 119934 (12); 15 km. ESE San Juan de la Punta, KU 23707; San Lorenzo, USNM 123508-12; 3 km. SW San Marcias KU 23841; 1.5 km. S Santa Rosa, UIMNH 42702; 2 km. S Santiago Tuxtla, UMMZ 121201 (4); Sauzel, UMMZ 121239; 14 km. E Suchil, UIMNH 46880; 15 km. S Tampico (Tamaulipas), UMMZ 103322 (4); 4 km. N Tapalapan, UMMZ 115186 (2); Tecolutla, UIMNH 42677-700; 16 km. NW Tehuatlan, UIMNH 3660-3; 5 km. S Tehuatlan, KU 23842; Teocelo, KU 26306; Tierra Colorado, CNHM 102393, 102395-6, UIMNH 30789-91; Veracruz, AMNH 6301-4, 59398-402, UIMNH 30801, UMMZ 115174, 122060 (2); 24 km. W Veracruz, CNHM 104570-2.

=Yucatan=: No specific locality, CNHM 548, 49067, USNM 32298; Chichen-Itza, CNHM 20636, 26938-49, 36559-62, UIMNH 30742-6, UMMZ 73173 (6), 73174 (14), 73175 (14), 73178-9, 76171, 83107 (2), 83108, 83109 (2), 83915 (30), USNM 72744; 9 km. E Chichen-Itza, KU 71438-9; 12 km. E Chichen-Itza, KU 71440; Merida, CNHM 40659-66, UIMNH 30747-8, UMMZ 73182; 6 km. S Merida, KU 75194; 8.8 km. SE Ticul, UMMZ 114296; Valladolid, CNHM 26934-6; Xcalah-op, CNHM 53906-14; 3.5 km. E Yokdzonot, KU 71441-3, 71720 (tadpoles).

BRITISH HONDURAS: Belize, CNHM 4153, 4384-5, 4387, UMMZ 75310, USNM 26065; Bokowina, CNHM 49064-5; Cocquercote, UMMZ 75331 (2); Cohune Ridge, UMMZ 80738 (15); Double Falls, CNHM 49066; El Cayo, UMMZ 75311; 6 km. S El Cayo, MCZ 37856; Gallon Jug, MCZ 37848-55; Manatee, CNHM 4264-7; Mountain Pine Ridge, MCZ 37857-8; San Augustin, UMMZ 80739; San Pedro, Columbia, MCZ 37860-2; Valentin, UMMZ 80735 (4), 80736 (2), 80737 (2); 5 km. S Waha Loaf Creek, MCZ 37859.

GUATEMALA: =Alta Verapaz=: 5.1 km. NE Campur, KU 68464 (tadpoles), 67465 (young); 28.3 km. NE Campur, KU 64203-22, 68183-4 (skeletons); Chama, MCZ 15792-3, UMMZ 90895 (7), 90896 (5), 90897 (29), 90898 (12), 90899; Chinaja, KU 55939-41, 57193-8, 60018-20 (tadpoles), 60021 (eggs), 60022 (tadpoles); Coban, CNHM 21006; Cubilquitz, UMMZ 90902 (10); Finca Canihor, UMMZ 90908; Finca Chicoyou, KU 57246-8, 60026 (young), 64202, 68466-7 (tadpoles); Finca Los Alpes, KU 64197-201, 68463 (tadpoles); Finca Los Pinales, UMMZ 90903 (2); Finca Tinajas, BYU 16031; Finca Volcan, UMMZ 90905 (4), 90906-7; Panzos, MNHN 6313, UMMZ 90904; Samac, UMMZ 90900; Samanzana, UMMZ 90901 (6).

=Baja Verapaz=: Chejel, UMMZ 90909 (7), 90910 (3); San Geronimo, UMMZ 84076 (16).

=Chiquimula=: 1.6 km. SE Chiquimula, UMMZ 98112; Esquipulas, UMMZ 106793 (28).

=El Peten=: 20 km. NNW Chinaja (Alta Verapaz), KU 57199-240; Flores, UMMZ 117985; La Libertad, KU 60024 (young), UMMZ 75313-20, 75323 (2), 75324 (7), 75325 (13), 75326 (2), 75327 (11), 75328 (12), 75329 (2); 3 km. SE La Libertad, KU 57243-4; 13 km. S La Libertad, MCZ 21458 (2); Pacomon, USNM 71334; Piedras Negras, USNM 114469-71; Poptun, UMMZ 120475; Poza de la Jicotea, USNM 114672; Ramate-Yaxha trail, UMMZ 75321; Rio de la Pasion between Sayaxche and Subin, KU 57151; Rio San Roman, 16 km. NNW Chinaja (Alta Verapaz), KU 55942-6; Sacluc, USNM 25131; Sayaxche, KU 57144-5; Tikal, UMMZ 117983 (7), 117984 (5), 117993 (5), 120474 (5); Toocog, KU 57241-2, 60023 (young), 60025 (young); Uaxactun, UMMZ 70401-3; Yaxha, UMMZ 75322; 19 km. E Yaxha, UMMZ 75330 (4).

=El Quiche=: Finca Tesoro, UMMZ 89166 (3), 90549 (tadpoles).

=Escuintla=: Rio Guacalate, Masagua, USNM 125239; Tiquisate, UMMZ 98262 (7).

=Guatemala=: 16 km. NE Guatemala, KU 43545-53.

=Huehuetenango=: Finca San Rafael, 16 km. SE Barillas, CNHM 40912-6; 45 km. WNW Huehuetenango, KU 64223-4; Jacaltenango, UMMZ 120080 (6), 120081 (14), 120082 (13).

=Izabal=: 2 km. SW Puerto Matias de Galvez, KU 60027-8 (tadpoles); Quirigua, CNHM 20587, UMMZ 70060.

=Jalapa=: Jalapa, UMMZ 98109, 106792 (11).

=Jutiapa=: Finca La Trinidad, UMMZ 107728 (10); Jutiapa, UMMZ 106789; 1.6 km. SE Mongoy, KU 43069; Santa Catarina Mita, UMMZ 106790.

=Progreso=: Finca Los Leones, UMMZ 106791.

=Quetzaltenango=: Coatepeque, AMNH 62204.

=Retalhueleu=: Casa Blanca, UMMZ 107725 (18); Champerico, UMMZ 107726 (3).

=San Marcos=: Talisman Bridge, USNM 128056-7.

=Santa Rosa=: Finca La Guardiana, UMMZ 107727 (6); Finca La Gloria, UMMZ 107724 (6); 1.6 km. WSW El Molino, KU 43065-8.

EL SALVADOR: =La Libertad=: 16 km. NW Santa Tecla, KU 43542-4. =Morazan=: Divisadero, USNM 73284. =San Salvador=: San Salvador, CNHM 65087-99, KU 61955-88, 62138-9 (skeletons), 62154 (eggs), 62155-60 (tadpoles), 68462 (tadpoles), UMMZ 117586 (3), 118380 (3), USNM 140278.

HONDURAS: State unknown: Guaimas, UMMZ 58391. =Atlantidad=: Isla de Roatan, CNHM 34551-4; La Ceiba, USNM 64985, 117589-91; Lancetilla, MCZ 16207-11; Tela, MCZ 15774-5, 28080, UMMZ 58418, USNM 82173-4. =Choluteca=: 1.5 km. NW Choluteca, KU 64228-32; 10 km. NW Choluteca, KU 64233; 10 km. E Choluteca, KU 64226-7; 12 km. E Choluteca, KU 64225; 5 km. S Choluteca, USC 2700 (2). =Colon=: Bambu, UF 320; Belfate, AMNH 45692-5; Patuca, USNM 20261. =Comayagua=: La Mision, 3.5 leagues N Siguatepeque, MCZ 26424-5. =Copan=: Copan, UMMZ 83026 (2). =Cortes=: Cofradia, AMNH 45345-6; Hacienda Santa Ana, CNHM 4724-31; Lago de Yojoa, MCZ 26410-1; Rio Lindo, AMNH 54972. =El Paraiso=: El Volcan, MCZ 26436. =Francisco Morazan=: Tegucigalapa, BYU 18301-4, 18837-41, MCZ 26395-7, USNM 60499. =Gracias A Dios=: Rio Segovia, MCZ 24543. =Santa Barbara=: Santa Barbara, USNM 128062-5.

NICARAGUA: =Chinandega=: 4 km. N, 2 km. W Chichigalpa, KU 85385; Chinandega, MCZ 2632; Rio Tama, USNM 40022; San Antonio, KU 84944-9 (skeletons), 85386-403. =Chontales=: 1 km. NE Acoyapa, KU 64234. =Esteli=: Finca Daraili, 5 km. N, 15 km. E Condega, KU 85404-8; Finca Venecia, 7 km. N, 16 km. E Condega, KU 85409. =Leon=: 1.6 km. ENE Poneloya, KU 43084-5. =Managua=: Managua, USNM 79989-90; 8 km. NW Managua, KU 43094-110; 20 km. NE Managua, KU 85412; 21 km. NE Managua, KU 85413-4; 5 km. SW Managua, KU 43086-93; 2 km. N Sabana Grande, KU 85411; 3 km. N Sabana Grande, KU 43070-8; 20 km. S, 0.5 km. W Tipitapa, KU 85410. =Matagalpa=: Guasqualie, UMMZ 116493; Matagalpa, UMMZ 116492; 19 km. N Matagalpa, UMMZ 116494. =Rio San Juan=: Greytown, USNM 19585-6, 19767-8. =Rivas=: Javillo, UMMZ 123001; Moyogalpa, Isla Ometepe, KU 85428-37, 87706 (tadpoles); Penas Blancas, KU 85417; Rio Javillo, 3 km. N, 4 km. W Sapoa, KU 85418-20, 85438 (skeleton); 13.1 km. SE Rivas, KU 85415; 14.8 km. SE Rivas, KU 85421-3; 11 km. S, 3 km. E Rivas, KU 85416; 16 km. S Rivas, MCZ 29009-10; 7.7 km. NE San Juan del Sur, KU 85426-7; 16.5 km. NE San Juan del Sur, KU 85424-5, 87705 (young); 5 km. SE San Pablo, KU 43079-83. =Zelaya=: Cooley, AMNM 7063-8, 8019-20, 8022, 8034-5; Cukra, AMNH 8016-7; Musahuas, Rio Huaspuc, AMNH 58428-31; 11 km. NW Rama, Rio Siquia, UMMZ 79708, 79709 (5), 79710 (2); Rio Escondido, USNM 19766, 20701; Rio Siquia at Rio Mico, UMMZ 79707 (10); Sioux Plantation, AMNH 7058-61, 8023-33.

COSTA RICA: =Alajuela=: Los Chiles, AMNH 54639; Orotina, MCZ 7960-1; San Carlos, USNM 29991. =Guanacaste=: La Cruz, USC 8232 (3); 4.3 km. NE La Cruz, UMMZ 123002; 18.4 km. S La Cruz, USC 8136; 23.5 km. S La Cruz, USC 8094 (4); 3 km. W La Cruz, USC 8233 (4); 2 km. NE Las Canas, KU 64235-7; Las Huecas, UMMZ 71212-3; Liberia, KU 36787, USC 8161; 11.5 km. N Liberia, USC 8149; 13 km. N Liberia, USC 8139; 22.4 km. N Liberia, USC 8126; 8 km. NNW Liberia, KU 64238; 8.6 km. ESE Playa del Coco, USC 8137; 21.8 km. ESE Playa del Coco, USC 8138; Rio Piedra, 1.6 km. W Bagaces, USC 7027; Rio Bebedero, 5 km. S Bebedero, KU 64158; 5 km. NE Tilaran, KU 36782-6. =Heredia=: 13 km. SW Puerto Viejo, KU 64142-6. =Limon=: Batan, KU 34927; Guacimo, USC 621; Pandora, USC 505 (3); Suretka, KU 36788-9; Tortugero, UF 7697, 10540-2. =Puntarenas=: Barranca, CNHM 35254-6; 15 km. WNW Barranca, KU 64155-7, UMMZ 118381; 18 km. WNW Barranca, UMMZ 118382 (4); 4 km. WNW Esparta, KU 64159-96, 68178-82 (skeletons); 19 km. NW Esparta, KU 64147-54.

=Smilisca cyanosticta= (Smith), new combination

_Hyla phaeota_, Taylor, Univ. Kansas Sci. Bull., 28(5):80, May 15,
1942. Taylor and Smith, Proc. U. S. Natl. Mus., 95(3185):589, June
30, 1945.

_Hyla phaeota cyanosticta_ Smith, Herpetologica, 8:150, Jan. 30,
1953 [Holotype.--USNM 111147 from Piedras Negras, El Peten,
Guatemala; Hobart M. Smith collector]. Shannon and Werler, Trans.
Kansas Acad. Sci., 58:386, Sept. 24, 1955. Poglayen and Smith,
Herpetologica, 14:11, April 25, 1958. Cochran, Bull. U. S. Natl.
Mus., 220:57, 1961. Smith, Illinois Biol. Mono., 32:25, May, 1964.

_Smilisca phaeota cyanosticta_, Stuart, Misc. Publ. Mus. Zool.
Univ. Michigan, 122:42, April 2, 1963. Duellman, Univ. Kansas Publ.
Mus. Nat. Hist., 15(5):229, Oct. 4, 1963.

_Diagnosis._--Size moderately large ([M] 56.0 mm., [F] 70.0 mm.); skull as long as wide; frontoparietal fontanelle large; narrow supraorbital flanges having irregular margins anteriorly; 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 usually more than 52 per cent of snout-vent length; labial stripe silvery-white; lips lacking vertical bars; loreal region pale green; pale bronze-colored stripe from nostril along edge of eyelid to point above tympanum narrow, bordered below by narrow dark brown stripe from nostril to eye, and broad dark brown postorbital mark encompassing tympanum and terminating above insertion of arm; flanks, dark brown with large pale blue spots; anterior and posterior surfaces of thighs dark brown with small pale blue spots on thighs. (Foregoing combination of characters distinguishing _S. cyanosticta_ from any other species in genus.)

_Description and Variation._--The largest males are from Piedras Negras, El Peten, Guatemala, and they average 52.5 mm. in snout-vent length whereas males from Los Tuxtlas, Veracruz, average 50.6 mm. and those from northern Oaxaca 50.3 mm. The smallest breeding male has a snout-vent length of 44.6 mm. The average ratio of tibia length to snout-vent length is 54.8 per cent in males from Piedras Negras, and 56.4 and 56.3 per cent in males from Los Tuxtlas and Oaxaca, respectively. The only other character showing noticeable geographic variation is the size of the tympanum. The average ratio of the diameter of the tympanum to the diameter of the eye is 76.3 per cent in males from Piedras Negras, 71.8 from Oaxaca, and 66.9 from Los Tuxtlas.

The dorsal ground color of _Smilisca cyanosticta_ is pale green to tan and the venter is creamy white. The dorsum is variously marked with dark olive-green or dark brown spots or blotches (Pl. 6B). An interorbital dark bar usually is present. The most extensive dark area is a V-shaped mark in the occipital region with the anterior branches not reaching the eyelids; this mark is continuous, by means of a narrow mid-dorsal mark, with an inverted V-shaped mark in the sacral region. In many specimens this dorsal marking is interrupted, resulting in irregular spots. In some specimens the dorsum is nearly uniform pale green or tan with a few small, dark spots. The hind limbs are marked by dark transverse bands, usually three or four each on the thigh and shank, and two or three on the tarsus. 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, which is bordered above by a narrow, bronze-colored stripe extending from the nostril along the edge of the eyelid to a point above the tympanum. The upper lip is white. A broad dark brown mark extends posteriorly from the orbit and encompasses the tympanum to a point above the insertion of the forelimb. The flanks are dark brown with many pale blue, rounded spots, giving the impression of a pale blue ground color with dark brown mottling enclosing spots. The anterior and posterior surfaces of the thighs are dark brown with many small pale blue spots. The inner surfaces of the shank and tarsus are colored like the posterior surfaces of the thighs. Pale blue spots are usually present on the proximal segments of the second and third toes. 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. The anal region is dark brown, bordered above by a narrow transverse white stripe. The throat in breeding males is dark, grayish brown with white flecks.

No geographic variation in the dorsal coloration is evident. Specimens from the eastern part of the range (Piedras Negras and Chinaja, Guatemala) have bold, dark reticulations on the flanks enclosing large pale blue or pale green spots, which fade to tan in preservative. Specimens from Oaxaca and Veracruz characteristically have finer dark reticulations on the flanks enclosing smaller blue spots; in many of these specimens the ventrolateral spots are smallest and are white.

All living adults are easily recognized by the presence of pale, usually blue, spots on the flanks and thighs. Individuals under cover by day have a tan dorsum with dark brown markings. A hiding individual at Chinaja, Alta Verapaz, Guatemala (KU 55936), had a pale tan dorsum when found; later the dorsal color changed to chocolate brown. A pale green patch was present below the eye; the spots on the posterior surfaces of the thighs were pale blue, and those on the flanks were yellowish green. A calling male obtained 10 kilometers north-northwest of Chinaja (KU 55934) was reddish brown when found at night; later the dorsal color changed to pale tan. A green patch below the eye was persistent. Usually these frogs are green at night. The coloration of an adult male (KU 87201) from 11 kilometers north of Vista Hermosa, Oaxaca, Mexico, was typical: "When calling dorsum pale green; later changed to dull olive-green. Flanks dark brown with pale blue spots in axilla and groin and bluish white flecks on mid-flank. Anterior and posterior surfaces of thighs, inner surfaces of shanks, and median dorsal surfaces of tarsi dark brown with blue spots. Canthal and postorbital stripes dark chocolate brown; labial stripe creamy white. Forearm, tarsal, and anal stripes pale cream-color. Throat dark brown with yellow flecks; belly and ventral surfaces of limbs creamy buff; webs pinkish tan; iris deep bronze, brown below pupil." (Duellman, field notes, June 24, 1964.)

Some individuals have both green and brown coloration in life. An individual obtained at night on the south slope of Volcan San Martin, Veracruz, Mexico, had a pale tan dorsum changing peripherally to pale green. The dorsal markings were dark brown and dark olive-green.

In contrast to the color-changes noted above, the labial region below the eye is always pale green, and pale spots are always present on the flanks and thighs in adults. The iris is invariably golden or bronze above and darker, usually brown, below. Minute black flecks are present on the iris, and in some individuals these flecks are so numerous that the eye appears gray.

Recently metamorphosed young have pale tan flanks, and the posterior surfaces of the thighs are orange-yellow; pale spots are absent. A juvenile (KU 55935) from Chinaja, Alta Verapaz, Guatemala, having a snout-vent length of 35.0 mm. was pale yellowish tan above with olive-green markings; the flanks were dark brown with pale blue spots, and the anterior and posterior surfaces of the thighs were uniform bright tomato red. A juvenile (UMMZ 121298), 18.6 mm. in snout-vent length, from the southeast slope of Volcan San Martin, Veracruz, Mexico, had pale tan flanks lacking blue spots, but had red thighs. Apparently the ontogenetic changes in coloration proceed as follows: (1) flanks pale tan and thighs orange-yellow, both lacking spots, (2) flanks pale tan and thighs red, both lacking spots, (3) flanks dark brown with blue spots and thighs red, lacking spots, and (4) flanks and thighs dark brown, both having pale blue spots.

_Natural History._--_Smilisca cyanosticta_ inhabits humid tropical forest and cloud forest from the lowlands to elevations of about 1200 meters in Los Tuxtlas and to about 900 meters in northern Oaxaca. In these moist environments the frogs apparently are active throughout the year. Active individuals have been obtained in January, July, and August in Los Tuxtlas, Veracruz, in June and July in northern Oaxaca, in February and March at Chinaja, Guatemala, and Taylor and Smith reported (1945:589) activity in May at Piedras Negras, Guatemala. Calling males were observed as follows; in a rain barrel 11 kilometers north of Vista Hermosa, Oaxaca, Mexico, on June 23-28, 1964; in a quiet pool in a stream 8 kilometers south of Yetla, Oaxaca, Mexico, in July, 1963 (Dale L. Hoyt, personal communication); in and near springs flowing into a stream at Dos Amates, Veracruz, Mexico, on August 4, 1959 (Douglas Robinson, personal communication); and in a water-filled depression in a log 10 kilometers west-northwest of Chinaja, Guatemala, on March 13, 1960. Taylor and Smith (1945:589) reported that individuals were found at night on the ground at the edge of temporary pools at Piedras Negras, Guatemala, on May 28-29, 1939. A clasping pair was found on a rock at the edge of a small stream on the south slope of Volcan San Martin, Veracruz, Mexico, on July 11, 1959 (Douglas Robinson, personal communication).

Only one individual has been observed in a tree at night. In the daytime, individuals were found in elephant ear plants (_Xanthosoma_) at Chinaja, Guatemala.

The breeding call consists of one or two moderately short notes that are lower pitched than those of _S. baudini_, but higher pitched than those of _S. phaeota_. Each note has a duration of 0.25 to 0.45 seconds and is repeated at intervals of one-half minute to several minutes. Each note is a vibrant "waunk," having 110 to 180 pulses per second and dominant frequency of 1600 to 2100 cycles per second (Pl. 10B).

Apparently the eggs are deposited as loose clumps in the water. In eggs in the yolk plug stage of development, the diameter of the embryo is about 2.3 mm.; that of the outer envelope is 4.0 mm. Hatchling tadpoles have total lengths of 5.8 to 6.5 mm. and body lengths of 2.8 to 3.1 mm. The external gills are moderately long, slender, and filamentous; the yolk sac is still moderately large. The body and anterior part of the caudal musculature are dark brown; posteriorly the caudal musculature is pale brown. The caudal fins are creamy tan. The oral discs are large and ovoid. The growth of the tadpole is summarized in Table 10.

A typical tadpole in stage 30 of development (KU 87652 from 11 km. N Vista Hermosa, Oaxaca, Mexico) can be described as follows:

Body length 9.5 mm.; tail length 15.5 mm.; total length 25.0 mm.; body slightly wider than deep; snout rounded laterally, broadly ovoid dorsally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, slightly posterior to midpoint of body and slightly below midline; anal tube dextral; caudal musculature slender, barely curved upward distally; dorsal fin not extending onto body, depth of dorsal fin slightly more than that of ventral fin on mid-length of tail; dorsal part of body dark brown; ventral surfaces transparent, lacking pigment; posterior edge of body pale cream-color; caudal musculature creamy white with interconnected brown spots; caudal fins transparent with small brown blotches on dorsal fin and posterior half of ventral fin; iris coppery bronze in life (Fig. 12). Mouth small, median part of upper lip bare; rest of mouth bordered by single row of bluntly rounded papillae; lateral fold present; tooth rows 2/3; all tooth-rows approximately equal in length; second upper row broadly interrupted medially; other rows complete; upper beak moderately deep, forming broad arch with slender lateral processes; lower beak slender, broadly V-shaped; both beaks finely serrate (Fig. 15C).

All tadpoles having fully developed mouth parts have 2/3 tooth rows. Little variation is noticeable in coloration. In many specimens the posterior edge of the body is dark brown instead of pale cream-color. Mottling is rather dense on the caudal fins in all specimens; in some individuals pigment is concentrated along the anterior one-third of the lateral groove. In life the body is dark brown with greenish gold flecks ventrally; the caudal musculature is gray.

In each of two recently metamorphosed young the snout-vent length is 14.0 mm. Coloration of young in life (KU 87653 from 11 km. N Vista Hermosa, Oaxaca, Mexico): "Dorsum pale tan with dark brown markings. Thighs orange-yellow; labial stripe white; iris bronze" (Duellman, field notes, July 10, 1964.)

_Remarks._--Smith (1953:150) named _cyanosticta_ as a subspecies of _Hyla phaeota_. The differences in cranial characters and certain external characters between _phaeota_ and _cyanosticta_ indicate that they are distinct species. Furthermore, a gap of about 350 kilometers exists between the known geographic ranges of the two kinds.

_Distribution._--_Smilisca cyanosticta_ inhabits wet forests on the Atlantic slope of southern Mexico and northern Central America from northern Oaxaca and southern Veracruz through northern Chiapas in Mexico and into El Peten and northern Alta Verapaz in Guatemala (Fig. 2). Apparently the range is discontinuous, for in southern Mexico the species is found in cloud forest at elevations of 830 to 900 meters on the northern slopes of the Sierra de Juarez. In the Sierra de Los Tuxtlas in southern Veracruz the species is found in wet forest at elevations of 300 to 1200 meters, but is absent in the intervening lowlands characterized by drier forest. In the west forests of northern Alta Verapaz and El Peten, Guatemala, the species is found at low elevations.

_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 Volcan San Martin, UMMZ 121200; Volcan San Martin, UIMNH 35403-4, 35408-12, UMMZ 118163; SE slope Volcan San Martin, UMMZ 121199, 121295 (2), 121296, 121298.

GUATEMALA: =Alta Verapaz=: Chinaja, KU 55935-7, 55938 (skeleton). =El=

=Peten=: 10 km. NNW Chinaja (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.

=Smilisca phaeota= (Cope)

_Hyla phaeota_ Cope, 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.
Muenchen, 27:215, 1897. Guenther, 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, Aug. 1, 1954. Goin,
Herpetologica, 14:120, July 23, 1958. Cochran, Bull. U. S. Natl.
Mus., 220:57, 1961.

_Hyla labialis_ Peters, Monats. Konigl. Akad. Wissen. Berlin, p.
463, 1863 [Holotype.--ZMB 4913 from "region of Bogota," 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 dolomedes_ Barbour, Occas. Papers Mus. Zool. Univ.
Michigan, 129:11, Jan. 25, 1923 [Holotype.--MCZ 8539 from Rio
Esnape, Sambu Valley, Darien, Panama; 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 distinguishing _S. phaeota_ from 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 on Ten Specimens From Each Locality.)

==================================================================
|Snout-vent |Head width/|Interorbital
Locality |length |snout-vent |distance/
| |length |head width
-----------------------------+-----------+-----------+------------
Bonanza, Nicaragua | 40.8-47.7 | 34.1-38.0 | 31.0-36.1
| (43.7) | (36.3) | (35.4)
-----------------------------+-----------+-----------+------------
Heredia Prov., Costa Rica | 46.3-59.0 | 32.5-36.0 | 30.5-39.6
| (51.7) | (35.0) | (34.7)
-----------------------------+-----------+-----------+------------
Puntarenas Prov., Costa Rica | 53.6-64.9 | 32.6-36.1 | 31.0-38.0
| (61.4) | (34.5) | (34.4)
-----------------------------+-----------+-----------+------------
Canal Zone, Panama | 52.4-65.5 | 33.5-37.6 | 31.3-37.2
| (56.5) | (35.6) | (34.7)
-----------------------------+-----------+-----------+------------
Rio Quesada, Colombia | 52.6-61.0 | 33.1-37.1 | 30.1-33.9
| (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 to snout-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 Panama, 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 of _Smilisca phaeota_ is 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 coloration from 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, Panama (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 phaeota_ inhabits 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, Panama, in February and in July, and Breder (1946:416) noted calling individuals in the Chucanaque drainage of Darien, Panama 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, Panama, 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 like _Smilisca 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 Rio Chitaria, 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, directed dorsolaterally; 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 Guapiles, 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 Rio Chitaria, 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) named _Hyla labialis_ from the "region of Bogota, Colombia", but in 1873 regarded his new species as identical with _Hyla phaeota_ Cope, 1862, from Turbo, Colombia. The name _Hyla labialis_ has 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 name _Hyla vilsoniana_ Cope, 1899, for the frogs from the northern Andes previously referred to _Hyla labialis_. A review of the nomenclature and taxonomy of these frogs, which superficially resemble _Smilisca_ but are unrelated, is beyond the scope of the present study.

_Hyla baudini dolomedes_ Barbour, 1923, is based on a small _Smilisca phaeota_ (MCZ 8539) having a snout-vent length of 45.5 mm. Dunn (1931a:413) placed _dolomedes_ in the synonymy of _Smilisca phaeota_. We have examined the holotype of _dolomedes_ and agree with Dunn's assignment.

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

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

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

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

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

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

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

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

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

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

PANAMA: No province: Cano Saddle, USNM 69588; Punta de Pena, USNM 38733; Quipo, AMNH 18925-6. =Bocas del Toro=: Almirante, KU 80080, 91835-6; 1.6 km. W Almirante, KU 91837; 3 km. W Almirante, KU 91824 (skeleton), 91838-43, 91906-7; 11 km. NW Almirante, CNHM 67853-61; 13 km. W Almirante, KU 91825-7 (skeletons), 91844-9; Fish Creek, KU 92329; Isla Popa, KU 91850-1. =Canal Zone=: Barro Colorado Island, CNHM 6007, 13316, 13325, 13331, 13360-2, 13377-8, MCZ 24191-5, UF 7523, UMMZ 63547-60, 64457, 69497 (3); 3.7 km. W Cocoli, KU 67916; Fort Sherman, MCZ 10139; Gatun, MCZ 35644; Junction roads C25B and C16, TNHC 23839; Madden Forest Preserve, TNHC 23837-8. =Cocle=: El Valle, KU 77521-4, 77649 (tadpole), TNHC 23369. =Comarca del Baru=: Progreso, UMMZ 61085-9. Colon: Achiote, KU 77516-20, 77648 (young); Rio Candelaria, CNHM 67851-2. =Darien=: Rio Esnape, Sambu Valley, MCZ 8539; Rio Sucubti, Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, Camp Townsend, AMNH 40988. =Panama=: NW slope Cerro Prominente, KU 80459; Finca La Sumbadora, KU 91914 (skeleton). =Chiriqui=: 2 km. W Concepcion, AMNH 68910.

COLUMBIA: =Antioquia=: Puerto Berrio, CNHM 30805 (Goin); Turbo, USNM 39899. =Caldas=: Pueblorrica, Santa Cecilia, CNHM 54768-71 (Goin). =Choco=: No specific locality, AMNH 3984-6; Andagoya, BMNH 1915.10.21. 69-70, CNHM 81857 (Goin); Golfo de Uraba, CNHM 63881 (Goin); Pena Lisa, Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro, CNHM 4451-3, 4455-61 (Goin); Rio San Juan, Playa del Oro, CNHM 54772 (Goin); Rio Quesada, AMNH 13615-77; 37 km. up Rio Pune, AMNH 13688; 48 km. up Rio Pune, AMNH 13689. =Narino=: Tumaco, Rio Rosario, CJG 2310-13 (Goin). =Valle=: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT 166, 346-47, 388 (Goin); Rio Calima above Cordoba, CJG 2249-57 (Goin).

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

=Smilisca puma= (Cope), new combination

_Hyla puma_ Cope, Proc. Amer. Philos. Soc., 22:183, 1885
[Holotype.--USNM 13735 from Nicaragua; J. F. Moser collector].
Guenther, Biologia Centrali-Americana: Reptilia and Batrachia, p.
270, Sept., 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 251,
June, 1923. Cochran, Bull. U. S. Natl. Mus., 220:58, 1961.

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

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

_Diagnosis._--Size small ([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 distinguishing _S. puma_ from other species in genus.)

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

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

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

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

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

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

Only six tadpoles are available for study. Four of them in stage 34 of development have body lengths of 9.0 to 9.5 mm., tail lengths of 14.0 to 15.0 mm., and total lengths of 23.0 to 24.5 mm. One tadpole in stage 38 and one in stage 40 have total lengths of 31.0 mm. A typical tadpole in stage 34 of development (KU 91807 from 7.5 km. W Puerto Viejo, Heredia Province, Costa Rica) has a body length of 9.5 mm., tail length of 15.0 mm., and total length of 24.5 mm.; body about three-fourths as deep as wide; snout rounded dorsally and laterally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, about two-thirds distance from snout to posterior end of body and slightly below midline; anal tube dextral; caudal musculature slender, barely curved upward distally; dorsal fin extending onto body; at mid-length of tail, depth of caudal musculature equal to that of dorsal fin and ventral fin; body grayish brown, palest ventrally; caudal musculature pale creamy yellow with bold gray reticulations; caudal fins transparent with gray reticulations anteriorly and black flecks posteriorly on both fins (Fig. 14A). Median part of upper lip bare; rest of mouth bordered by two rows of short blunt papillae; lateral fold present; tooth-rows 2/3; upper rows equal in length; second upper row broadly interrupted medially; three lower rows complete, first and second rows equal in length, slightly shorter than upper rows; third lower row noticeably shorter; upper beak shallow, forming broad, continuous arch with slender lateral processes; lower beak slender, broadly V-shaped, both beaks finely serrate (Fig. 15B).

All six tadpoles are colored alike, except that in the larger specimens scattered white flecks are present on the ventral surface of the body, and the dark reticulations continue farther posteriorly on the caudal fins than in the smaller tadpoles. In two specimens the third lower tooth-row is only about one-half the length of the other lower rows, and in one specimen the second lower tooth-row is shorter than the first. Coloration of tadpoles in life: "Body olive-brown with silvery green flecks laterally. Caudal musculature olive-brown with greenish tan flecks. Fins brown with greenish gold flecks. Iris deep bronze." (Duellman, field notes, February 19, 1965).

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