Neotropical Hylid Frogs, Genus Smilisca Part 4

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 f.l.a.n.g.es 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; l.a.b.i.al 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 pos...o...b..tal mark encompa.s.sing 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 distinguis.h.i.+ng _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 encompa.s.ses 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 a.n.a.l 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 pos...o...b..tal stripes dark chocolate brown; l.a.b.i.al stripe creamy white. Forearm, tarsal, and a.n.a.l 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 l.a.b.i.al 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; a.n.a.l 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; l.a.b.i.al 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.