University of Kansas Publications
Museum of Natural History

Neotropical Hylid Frogs, Genus Smilisca

BY

WILLIAM E. DUELLMAN AND LINDA TRUEB

University of Kansas Lawrence

University of Kansas Publications, Museum of Natural History

University of Kansas

PRINTED BY
ROBERT R. (BOB) SANDERS, STATE PRINTER
TOPEKA, KANSAS
1966

Neotropical Hylid Frogs, Genus Smilisca

BY

WILLIAM E. DUELLMAN AND LINDA TRUEB

CONTENTS

INTRODUCTION

The family Hylidae, as currently recognized, is composed of about 34 genera and more than 400 species. Most genera (30) and about 350 species live in the American tropics. Hyla and 10 other genera inhabit Central America; four of those 10 genera (Gastrotheca, Hemiphractus, Phrynohyas, and Phyllomedusa) are widely distributed in South America. The other six genera are either restricted to Central America or have their greatest differentiation there. Plectrohyla and Ptychohyla inhabit streams in the highlands of southern Mexico and northern Central America; Diaglena and Triprion are casque-headed inhabitants of arid regions in México and northern Central America. Anotheca is a tree-hole breeder in cloud forests in Middle America. The genus Smilisca is the most widespread geographically and diverse ecologically of the Central American genera.

The definition of genera in the family Hylidae is difficult owing to the vast array of species, most of which are poorly known as regards their osteology, colors in life, and modes of life history. The genera Diaglena, Triprion, Tetraprion, Osteocephalus, Trachycephalus, Aparasphenodon, Corythomantis, Hemiphractus, Pternohyla, and Anotheca have been recognized as distinct from one another and from the genus Hyla on the basis of various modifications of dermal bones of the cranium. Phyllomedusa is recognized on the basis of a vertical pupil and opposable thumb; Plectrohyla is characterized by the presence of a bony prepollex and the absence of a quadratojugal. Gastrotheca is distinguished from other hylids by the presence of a pouch in the back of females. A pair of lateral vocal sacs behind the angles of the jaws and the well-developed dermal glands were used by Duellman (1956) to distinguish Phrynohyas from Hyla. He (1963a) cited the ventrolateral glands in breeding males as diagnostic of Ptychohyla. Some species groups within the vaguely defined genus Hyla have equally distinctive characters. The Hyla septentrionalis group is characterized by a casque-head, not much different from that in the genus Osteocephalus (Trueb, MS). Males in the Hyla maxima group have a protruding bony prepollex like that characteristically found in Plectrohyla.

Ontogenetic development, osteology, breeding call, behavior, and ecology are important in the recognition of species. By utilizing the combination of many morphological and biological factors, the genus Smilisca can be defined reasonably well as a natural, phyletic assemblage of species. Because the wealth of data pertaining to the morphology and biology of Smilisca is lacking for most other tree frogs in Middle America it is not possible at present to compare Smilisca with related groups in more than a general way.

Smilisca is an excellent example of an Autochthonous Middle American genus. As defined by Stuart (1950) the Autochthonous Middle American fauna originated from "hanging relicts" left in Central America by the ancestral fauna that moved into South America and differentiated there at a time when South America was isolated from North and Middle America. The genus Smilisca, as we define it, consists of six species, all of which occur in Central America. One species ranges northward to southern Texas, and one extends southward on the Pacific lowlands of South America to Ecuador. We consider the genus Smilisca to be composed of rather generalized hylids. Consequently, an understanding of the systematics and zoogeography of the genus can be expected to be of aid in studying more specialized members of the family.

Acknowledgments

Examination of many of the specimens used in our study was possible only because of the cooperation of the curators of many systematic collections. For lending specimens or providing working space in their respective institutions we are grateful to Doris M. Cochran, Alice G. C. Grandison, Jean Guibe, Robert F. Inger, Günther Peters, Gerald Raun, William J. Riemer, Jay M. Savage, Hobart M. Smith, Wilmer W. Tanner, Charles F. Walker, Ernest E. Williams, and Richard G. Zweifel.

We are indebted to Charles J. Cole and Charles W. Myers for able assistance in the field. The cooperation of Martin H. Moynihan at Barro Colorado Island, Charles M. Keenan of Corozal, Canal Zone, and Robert Hunter of San José, Costa Rica, is gratefully acknowledged. Jay M. Savage turned over to us many Costa Rican specimens and aided greatly in our work in Costa Rica. James A. Peters helped us locate sites of collections in Ecuador and Coleman J. Goin provided a list of localities for the genus in Colombia.

We especially thank Charles J. Cole for contributing the information on the chromosomes, and Robert R. Patterson for preparing osteological specimens. We thank M. J. Fouquette, Jr., who read the section on breeding calls and offered constructive criticism.

Permits for collecting were generously provided by Ing. Rodolfo Hernandez Corzo in México, Sr. Jorge A. Ibarra in Guatemala, and Ing. Milton Lopez in Costa Rica. This report was made possible by support from the National Science Foundation (Grants G-9827 and GB-1441) and the cooperation of the Museum of Natural History at the University of Kansas. Some of the field studies were carried out in Panamá under the auspices of a grant from the National Institutes of Health (NIH GM-12020) in cooperation with the Gorgas Memorial Laboratory in Panamá.

Materials and Methods

In our study we examined 4151 preserved frogs, 93 skeletal preparations, 88 lots of tadpoles and young, and six lots of eggs. We have collected specimens in the field of all of the species. Observations on behavior and life history were begun by the senior author in México in 1956 and completed by us in Central America in 1964 and 1965.

Osteological data were obtained from dried skeletons and cleaned and stained specimens of all species, plus serial sections of the skull of Smilisca baudini. Developmental stages to which tadpoles are assigned are in accordance with the table of development published by Gosner (1960). Breeding calls were recorded in the field on tape using Magnemite and Uher portable tape recorders. Audiospectrographs were made by means of a Vibralyzer (Kay Electric Company). External morphological features were measured in the manner described by Duellman (1956). In the accounts of the species we have attempted to give a complete synonymy. At the end of each species account the localities from which specimens were examined are listed alphabetically within each state, province, or department, which in turn are listed alphabetically within each country. The countries are arranged from north to south. Abbreviations for museum specimens are listed below:

Genus Smilisca Cope, 1865

Smilisca Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct., 1865 [Type species Smilisca daulinia Cope, 1865 = Hyla baudini Duméril and Bibron, 1841]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:75, June 17, 1948. Starrett, Copeia, 4:300, December 30, 1960. Goin, Ann. Carnegie Museum, 36:15, July 14, 1961.

Definition.—Medium to large tree frogs having: (1) broad, well ossified skull (consisting of a minimum amount of cartilage and/or secondarily ossified cartilage), (2) no dermal co-ossification, (3) quadratojugal and internasal septum present, (4) large ethmoid, (5) M. depressor mandibulae consisting of two parts, one arising from dorsal fascia and other from posterior arm of squamosal, (6) divided M. adductor mandibulae, (7) paired subgular vocal sacs in males, (8) no dermal appendages, (9) pupil horizontally elliptical (10) small amounts of amines and other active substances in skin, (11) chromosome number of N = 12 and 2N = 24, (12) breeding call consisting of poorly modulated, explosive notes, and (13) 2/3 tooth-rows in tadpoles.

Composition of genus.—As defined here the genus Smilisca contains six recognizable species. An alphabetical list of the specific and subspecific names that we consider to be applicable to species of Smilisca recognized herein is given below.

Distribution of genus.—Most of lowlands of México and Central America, in some places to elevations of nearly 2000 meters, southward from southern Sonora and Río Grande Embayment of Texas, including such continental islands as Isla Cozumel, México, and Isla Popa and Isla Cebaco, Panamá, to northern South America, where known from Caribbean coastal regions and valleys of Río Cauca and Río Magdalena in Colombia, and Pacific slopes of Colombia and northern Ecuador.

Key to Adults

Key to Tadpoles

ACCOUNTS OF SPECIES

Smilisca baudini (Duméril and Bibron)

Hyla baudini Duméril and Bibron, Erpétologie général, 8:564, 1841 [Holotype.—MNHN 4798 from "Mexico;" Baudin collector]. Günther, Catalogue Batrachia Salientia in British Museum, p. 105, 1858. Brocchi, Mission scientifique au Mexique ..., pt. 3, sec. 2, Études sur les batrachiens, p. 29, 1881. Boulenger, Catalogue Batrachia Salientia in British Museum, p. 371, Feb. 1, 1882. Werner, Abhand. Zool.-Bot. Gesell. Wien., 46:8, Sept. 30, 1896. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p. 270, Sept. 1901. Werner, Abhand. Konigl. Akad. Wiss. Munchen, 22:351, 1903. Cole and Barbour, Bull. Mus. Comp. Zool., 50(5):154, Nov. 1906. Gadow, Through southern México, p. 76, 1908. Ruthven, Zool. Jahr. 32(4):310, 1912. Decker, Zoologica, 2:12, Oct., 1915. Stejneger and Barbour, A checklist of North American amphibians and reptiles, p. 32, 1917. Noble, Bull. Amer. Mus. Nat. Hist., 38(10):341, June 20, 1918. Nieden, Das Tierreich, Amphibia, Anura I, p. 243, June, 1923. Gadow, Jorullo, p. 54, 1930. Dunn and Emlen, Proc. Acad. Nat. Sci. Philadelphia, 84:24, March 22, 1932. Kellogg, Bull. U. S. Natl. Mus., 160:160, March 31, 1932. Martin, Aquarien Berlin, p. 92, 1933. Stuart, Occas. Papers Mus. Zool., Univ. Michigan, 292:7, June 29, 1934; Misc. Publ. Mus. Zool. Univ. Michigan, 29:38, Oct. 1, 1935. Gaige, Carnegie Inst. Washington, 457:293, Feb. 5, 1936. Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 360:5, Nov. 20, 1937. Smith, Occas. Papers Mus. Zool. Univ. Michigan, 388:2, 12, Oct. 31, 1938; Ann. Carnegie Mus., 27:312, March 14, 1939. Taylor, Copeia, 2:98, July 12, 1939. Hartweg and Oliver, Misc. Publ. Mus. Zool. Univ. Michigan, 47:12, July 13, 1940. Schmidt and Stuart, Zool. Ser. Field Mus. Nat. Hist., 24(21):238, August 30, 1941. Schmidt, Zool. Ser. Field Mus. Nat. Hist., 22(8):486, Dec. 30, 1941. Wright and Wright, Handbook of frogs and toads, Ed. 2, p. 134, 1942. Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 471:15, May 17, 1943. Bogert and Oliver, Bull. Amer. Mus. Nat. Hist., 83(6):343, March 30, 1945. Taylor and Smith, Proc. U. S. Natl. Mus., 95(3185): 590, June 30, 1945. Smith, Ward's Nat. Sci. Bull., 1, p. 3, Sept., 1945. Schmidt and Shannon, Fieldiana, Zool. Chicago Nat. Hist. Mus., 31(9):67, Feb. 20, 1947. Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:26, June 12, 1948. Wright and Wright, Handbook of frogs and toads, Ed. 3, p. 298, 1949. Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 45:22, May, 1950. Mertens, Senckenbergiana, 33:170, June 15, 1952; Abhand. Senckenb. Naturf. Gesell., 487:28, Dec. 1, 1952. Schmidt, A checklist of North American amphibians and reptiles, Ed. 6, p. 69, 1953. Stuart Contr. Lab. Vert. Biol. Univ. Michigan, 68:46, Nov. 1954. Zweifel and Norris, Amer. Midl. Nat., 54(1):232, July 1955. Martin, Amer. Nat., 89:356, Dec. 1955. Duellman, Copeia, 1:49, Feb. 21, 1958. Goin, Herpetologica, 14:119, July 23, 1958. Turner, Herpetologica, 14:192, Dec. 1, 1958. Conant, A field guide to reptiles and amphibians, p. 284, 1958. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 13(2):59, Aug. 16, 1960; Univ. Kansas Publ., Mus. Nat. Hist., 15(1): 46, Dec. 20, 1961. Porter, Herpetologica, 18:165, Oct. 17, 1962.

Hyla vanvlietii Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:61, April 27, 1854 [Holotype.—USNM 3256 from Brownsville, Cameron County, Texas; S. Van Vliet collector]. Baird, United States and Mexican boundary survey, 2:29, 1859. Smith and Taylor, Univ. Kansas Sci. Bull., 33:361, March 20, 1950. Cochran, Bull. U. S. Natl. Mus., 220:60, 1961.

Hyla vociferans Baird, United States and Mexican boundary survey, 2:35 1859 [nomen nudum]. Diáz de León, Indice de los batracios que se encuentran en la República Mexicana, p. 20, June 1904.

Hyla muricolor Cope, Proc. Acad. Nat. Sci. Philadelphia, 14(9):359, 1862 [Holotype.—USNM 25097 from Mirador, Veracruz, México; Charles Sartorius collector]. Smith and Taylor, Univ. Kansas Sci. Bull., 33:349, March 20, 1950. Cochran, Bull. U. S. Natl. Mus., 220:56, 1961.

Smilisca daulinia Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct. 1865 [Holotype.—"skeleton in private anatomical museum of Hyrtl, Professor of Anatomy in the University of Vienna">[. Smith and Taylor, Univ. Kansas Sci. Bull., 33:347, March 20, 1950.

Smilisca daudinii [lapsus for baudini], Cope, Proc. Acad. Nat. Sci. Philadelphia, 23, pt. 2:205, 1871.

Smilisca baudini, Cope, Bull. U. S. Nat. Mus., 1:31, 1875; Jour. Acad. Nat. Sci. Philadelphia, 8, pt. 2:107, 1876; Proc. Amer. Philos. Soc., 18:267, August 11, 1879. Yarrow, Bull. U. S. Nat. Mus., 24:176, July 1, 1882. Cope, Bull. U. S. Nat. Mus., 32:13, 1887; Bull. U. S. Nat. Mus., 34:379, April 9, 1889. Dickerson, The frog book, p. 151, July, 1906. Smith and Taylor, Univ. Kansas Sci. Bull., 33:442, March 20, 1950; Taylor, U. Kan. Sc. Bull., 34:802, Feb. 15, 1952; Univ. Kansas Sci. Bull., 35:794, July 1, 1952. Brattstrom, Herpetologica, 8(3):59, Nov. 1, 1952. Taylor, U. Kan. Sci. Bull., 35:1592, Sept. 10, 1953. Peters, Occas. Papers Mus. Zool. Univ. Michigan, 554:7, June 23, 1954. Duellman, Occas. Papers Mus. Zool. Univ. Michigan, 560:8, Oct. 22, 1954. Chrapliwy and Fugler, Herpetologica, 11:122, July 15, 1955. Smith and Van Gelder, Herpetologica, 11:145, July 15, 1955. Lewis and Johnson, Herpetologica, 11:178, Nov. 30, 1955. Martin, Misc. Publ. Mus. Zool. Univ. Michigan, 101:53, April 15, 1958. Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 75:17, June, 1958. Minton and Smith, Herpetologica, 17:74, July 11, 1961. Nelson and Hoyt, Herpetologica, 17:216, Oct. 9, 1961. Holman, Copeia, 2:256, July 20, 1962. Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 122:41, April 2, 1963. Maslin, Herpetologica, 19:124, July 3, 1963. Holman and Birkenholz, Herpetologica, 19:144, July 3, 1963. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15(5):228, Oct. 4, 1963. Zweifel, Copeia, 1:206, March 26, 1964. Duellman and Klaas, Copeia, 2:313, June 30, 1964. Davis and Dixon, Herpetologica, 20:225, January 25, 1965. Neill, Bull. Florida State Mus., 9:89, April 9, 1965.

Hyla pansosana Brocchi, Bull. Soc. Philom., ser. 7, 1:125, 1877 [Holotype.—MNHN 6313 from Panzós, Alta Verapaz, Guatemala; M. Bocourt collector]; Mission scientifique au Mexique ..., pt. 3, sec. 2, Études sur les batrachiens, p. 34, 1881.

Hyla baudini baudini, Stejneger and Barbour, A checklist of North American amphibians and reptiles, Ed. 3, p. 34, 1933. Wright and Wright, Handbook of frogs and toads, p. 110, 1933. Stejneger and Barbour, A checklist of North American amphibians and reptiles, Ed. 4, p. 39, 1939; A checklist of North American amphibians and reptiles, Ed. 5, p. 49, 1943. Smith and Laufe, Trans. Kansas Acad. Sci., 48(3):328, Dec. 19, 1945. Peters, Nat. Hist. Misc., 143:7, March 28, 1955.

Hyla beltrani Taylor, Univ. Kansas Sci. Bull. 28(14):306, Nov. 15, 1942 [Holotype.—UIMNH 25046 (formerly EHT-HMS 29563) from Tapachula, Chiapas, México; A. Magaña collector]. Smith and Taylor, Bull. U. S. Natl. Mus. 194:87, June 17, 1948; Univ. Kansas Sci. Bull, 33:326, March 20, 1950. Smith, Illinois Biol. Mono., 32:23, May, 1964.

Smilisca baudini baudini, Smith, Jour. Washington Acad. Sci., 37(11):408, Nov. 15, 1947. Smith and Taylor, Bull. U. S. Natl. Mus., 194:75, June 17, 1948; Univ. Kansas Sci. Bull., 33:347, March 20, 1950. Brown, Baylor Univ. Studies, p. 68, 1950. Smith, Smith, and Werler, Texas Jour. Sci., 4(2):254, June 30, 1952. Smith and Smith, Anales Inst. Biol., 22(2):561, Aug. 7, 1952. Smith and Darling, Herpetologica, 8(3):82, Nov. 1, 1952. Davis and Smith, Herpetologica, 8(4):148, Jan. 30, 1953. Neill and Allen, Publ. Res. Div. Ross Allen's Reptile Inst., 2(1):26, Nov. 10, 1959. Maslin, Univ. Colorado Studies, Biol. Series, 9:4, Feb. 1963. Holman, Herpetologica, 20:48, April 17, 1964.

Hyla manisorum Taylor, Univ. Kansas Sci. Bull., 36:630, June 1, 1954 [Holotype.—KU 34927 from Batán, Limón Province, Costa Rica; Edward H. Taylor collector]. Duellman and Berg, Univ. Kansas Publ. Mus. Nat. Hist, 15(4):193, Oct. 26, 1962.

Diagnosis.—Size large ([M] 76 mm., [F] 90 mm.); skull noticeably wider than long, having small frontoparietal fontanelle (roofed with bone in large individuals); postorbital processes long, pointed, curving along posterior border of orbit; squamosal large, contacting maxillary; tarsal fold strong, full length of tarsus; inner metatarsal tubercle large, high, elliptical; hind limbs relatively short, tibia length less than 55 per cent snout-vent length; lips strongly barred with brown and creamy tan; flanks pale cream with bold brown or black reticulations in groin; posterior surfaces of thighs brown with cream-colored flecks; dorsal surfaces of limbs marked with dark brown transverse bands. (Foregoing combination of characters distinguishing S. baudini from any other species in genus.)

Description and Variation.—Considerable variation in size, and in certain proportions and structural characters was observed; variation in some characters seems to show geographic trends, whereas variation in other characters apparently is random. Noticeable variation is evident in coloration, but this will be discussed later.

In order to analyze geographic variation in size and proportions, ten adult males from each of 14 samples from various localities throughout the range of the species were measured. Snout-vent length, length of the tibia in relation to snout-vent length, and relative size of the tympanum to the eye are the only measurements and proportions that vary noticeably (Table 1). The largest specimens are from southern Sinaloa; individuals from the Atlantic lowlands of Alta Verapaz in Guatemala, Honduras, and Costa Rica are somewhat smaller, and most specimens from the Pacific lowlands of Central America are slightly smaller than those from the Atlantic lowlands. The smallest males are from the Atlantic lowlands of México, including Tamaulipas, Veracruz, the Yucatán Peninsula, and British Honduras.

The ratio of the tibia to the snout-vent length varies from 42.1 to 53.6 in the 14 samples analyzed. The average ratio in samples from the Pacific lowlands varies from 44.9 in Sinaloa and El Salvador to 47.8 in Guerrero; on the Gulf lowlands of México the average ratio varies from 45.6 in Veracruz to 47.6 on Isla del Carmen, Campeche. Specimens from the Yucatán Peninsula and the Caribbean lowlands have relatively longer legs; the variation in average ratios ranges from 49.1 in British Honduras to 51.2 in Costa Rica and 51.5 in Honduras.

Specimens from southern Sinaloa are outstanding in the large size of the tympanum; the tympanum/eye ratio varies from 84.2 to 94.4 (average 87.8). In most other samples the variation in average ratios ranges from 72.2 to 79.3, but specimens from Veracruz have an average ratio of 70.0; Campeche, 65.7; Honduras, 67.0; and Limón, Costa Rica, 68.5.

No noticeable geographic trends in size and proportions are evident. Specimens from southern Sinaloa are extreme in their large size, relatively short tibia, and large tympani, but in size and relative length of the tibia the Sinaloan frogs are approached by specimens from such far-removed localities as San Salvador, El Salvador, and Chinajá, Guatemala. Frogs from the Caribbean lowlands of Honduras and Costa Rica are relatively large and have relatively long tibiae and small tympani.

The inner metatarsal tubercle is large and high and its shape varies. The tubercle is most pronounced in specimens from northwestern México, Tamaulipas, and the Pacific lowlands of Central America. Possibly the large tubercle is associated with drier habitats, where perhaps the frogs use the tubercles for digging.

The ground color of Smilisca baudini is pale green to brown dorsally and white to creamy yellow ventrally. The dorsum is variously marked with dark brown or dark olive-green spots or blotches (Pl. 6A). In most specimens a dark interorbital bar extends across the head to the lateral edges of the eyelid; usually this bar is connected medially to a large dorsal blotch. There is no tendency for the markings on the dorsum to form transverse bands or longitudinal bars. In specimens from the southern part of the range the dorsal dark markings are often fragmented into small spots, especially posteriorly. The limbs are marked by dark transverse bands, usually three on the forearm, three on the thigh, and three or four on the shank. Transverse bands also are present on the tarsi and proximal segments of the fingers and toes. The webbing on the hands and feet is pale grayish brown. The loreal region and upper lip are pale green or tan; the lip usually is boldly marked with broad vertical dark brown bars, especially evident is the bar beneath the eye. A dark brown or black mark extends from the tympanum to a point above the insertion of the forearm; in some specimens this black mark is narrow or indistinct, but in most individuals it is quite evident. The flanks are pale gray to creamy white with brown or black mottling, which sometimes forms reticulations enclosing white spots. The anterior surfaces of the thighs usually are creamy white with brown mottling, whereas the posterior surfaces of the thighs usually are brown with small cream-colored flecks. A distinct creamy white anal stripe usually is present. Usually, there are no white stripes on the outer edges of the tarsi and forearms. In breeding males the throat is gray.

Most variation in coloration does not seem to be correlated with geography. The lips are strongly barred in specimens from throughout the range of the species, except that in some specimens from southern Nicaragua and Costa Rica the lips are pale and in some specimens the vertical bars are indistinct. Six specimens from 7.3 kilometers southwest of Matatán, Sinaloa, are distinctively marked. The dorsum is uniformly grayish green with the only dorsal marks being on the tarsi; canthal and post-tympanic dark marks absent. A broad white labial stripe is present and interrupted by a single vertical dark mark below the eye. A white stripe is present on the outer edge of the foot. The flanks and posterior surfaces of the thighs are creamy white, boldly marked with black. Two specimens from Alta Verapaz, Guatemala (CNHM 21006 from Cobán and UMMZ 90908 from Finca Canihor), are distinctive in having many narrow transverse bands on the limbs and fine reticulations on the flanks. Two specimens from Limón Province, Costa Rica (KU 34927 from Batán and 36789 from Suretka), lack a dorsal pattern; instead these specimens are nearly uniform brown above and have only a few small dark brown spots on the back and lack transverse bands on the limbs. The post-tympanic dark marks and dark mottling on the flanks are absent. Specimens lacking the usual dorsal markings are known from scattered localities on the Caribbean lowlands from Guatemala to Costa Rica.

The coloration in life is highly variable; much of the apparent variation is due to metachrosis, for individuals of Smilisca baudini are capable of undergoing drastic and rapid change in coloration. When active at night the frogs usually are pale bright green with olive-green markings, olive-green with brown markings, or pale brown with dark brown markings. The dark markings on the back and dorsal surfaces of the limbs are narrowly outlined by black. The pale area below the eye and just posterior to the broad suborbital dark bar is creamy white, pale green, or ashy gray in life. The presence of this mark is an excellent character by which to identify juveniles of the species. The flanks are creamy yellow, or yellow with brown or black mottling. In most individuals the belly is white, but in specimens from southern El Petén and northern Alta Verapaz, Guatemala, the belly is yellow, especially posteriorly. The iris varies from golden bronze to dull bronze with black reticulations, somewhat darker ventrally.

Natural History.—Throughout most of its range Smilisca baudini occurs in sub-humid habitats; consequently the activity is controlled by the seasonal nature of the rainfall and usually extends from May or June through September. Throughout México and Central America the species is known to call and breed in June, July, and August. Several records indicate that the breeding season in Central America is more lengthy. Gaige, Hartweg, and Stuart (1937:4) noted gravid females collected at El Recreo, Nicaragua, in August and September. Schmidt (1941:486) reported calling males in February in British Honduras. Stuart (1958:17) stated that tadpoles were found in mid-February, juveniles in February and March and half-grown individuals from mid-March to mid-May at Tikal, El Petén, Guatemala. Stuart (1961:74) reported juveniles from Tikal in July, and that individuals were active at night when there had been light rain in the dry season in February and March in El Petén, Guatemala. Smilisca baudini seeks daytime retreats in bromeliads, elephant-ear plants (Xanthosoma), and beneath bark or in holes in trees. By far the most utilized retreat in the dry season in parts of the range is beneath the outer sheaths of banana plants. Large numbers of these frogs were found in banana plants at Cuautlapan, Veracruz, in March, 1956, in March and December, 1959.

Large breeding congregations of this frog are often found at the time of the first heavy rains in the wet season. Gadow (1908:76) estimated 45,000 frogs at one breeding site in Veracruz. In the vicinity of Tehuantepec, Oaxaca, large numbers of individuals were found around rain pools and roadside ditches in July, 1956, and July, 1958; large concentrations were found near Chinajá, Guatemala, in June, 1960, and near Esparta, Costa Rica in July, 1961. Usually males call from the ground at the edge of the water or not infrequently sit in shallow water, but sometimes males call from bushes and low trees around the water. Stuart (1935:38) recorded individuals calling and breeding throughout the day at La Libertad, Guatemala. Smilisca baudini usually is absent from breeding congregations of hylids; frequently S. baudini breeds alone in small temporary pools separated from large ponds where numerous other species are breeding. In Guerrero and Oaxaca, México, S. baudini breeds in the same ponds with Rhinophrynus dorsalis, Bufo marmoreus, Engystomops pustulosus, and Diaglena reticulata, and in the vicinity of Esparta, Costa Rica, S. baudini breeds in ponds with Bufo coccifer, Hyla staufferi, and Phrynohyas venulosa. In nearly all instances the breeding sites of S. baudini are shallow, temporary pools.

The breeding call of Smilisca baudini consists of a series of short explosive notes. Each note has a duration of 0.09 to 0.13 seconds; two to 15 notes make up a call group. Individual call groups are spaced from about 15 seconds to several minutes apart. The notes are moderately high-pitched and resemble "wonk-wonk-wonk." Little vibration is discernible in the notes, which have 140 to 195 pulses per second and a dominant frequency of 2400 to 2725 cycles per second (Pl. 10A).

The eggs are laid as a surface film on the water in temporary pools. The only membrane enclosing the individual eggs is the vitelline membrane. In ten eggs (KU 62154 from San Salvador, El Salvador) the average diameter of the embryos in first cleavage is 1.3 mm. and of the vitelline membranes, 1.5 mm. Hatchling tadpoles have body lengths of 2.6 to 2.7 mm. and total lengths of 5.1 to 5.4 mm. The body and caudal musculature is brown; the fins are densely flecked with brown. The gills are long and filamentous. Growth and development of tadpoles are summarized in Table 9.

A typical tadpole in stage 30 of development (KU 60018 from Chinajá, Alta Verapaz, Guatemala) has a body length of 8.7 mm., a tail length of 13.6 mm., and a total length of 22.3 mm.; body slightly wider than deep; snout rounded dorsally and laterally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, located about midway on length of body and slightly below midline; anal tube dextral; caudal musculature slender, slightly curved upward distally; dorsal fin extending onto body, deepest at about one-third length of tail; depth of dorsal fin slightly more than that of ventral fin at mid-length of tail; dorsal part of body dark brown; pale crescent-shaped mark on posterior part of body; ventral surfaces transparent with scattered brown pigment ventrolaterally, especially below eye; caudal musculature pale tan with a dark brown longitudinal streak on middle of anterior one-third of tail; dorsum of anterior one-third of tail dark brown; brown flecks and blotches on rest of caudal musculature, on all of dorsal fin, and on posterior two-thirds of ventral fin; iris bronze in life (Fig. 11). Mouth small; median third of upper lip bare; rest of mouth bordered by two rows of conical papillae; lateral fold present; tooth rows 2/3; two upper rows about equal in length; second row broadly interrupted medially, three lower rows complete, first and second equal in length, slightly shorter than upper rows; third lower row shortest; first upper row sharply curved anteriorly in midline; upper beak moderately deep, forming a board arch with slender lateral processes; lower beak more slender, broadly V-shaped; both beaks bearing blunt serrations (Fig. 15A).

In tadpoles having fully developed mouthparts the tooth-row formula of 2/3 is invariable, but the coloration is highly variable. The color and pattern described above is about average. Some tadpoles are much darker, such as those from 11 kilometers north of Vista Hermosa, Oaxaca, (KU 87639-44), 3.5 kilometers east of Yokdzonot, Yucatán (KU 71720), and 4 kilometers west-southwest Puerto Juárez, Quintana Roo, México (KU 71721), whereas others, notably from 17 kilometers northeast of Juchatengo, Oaxaca, México (KU 87645), are much paler and lack the dark markings on the caudal musculature. The variation in intensity of pigmentation possibly can be correlated with environmental conditions, especially the amount of light. In general, tadpoles that were found in open, sunlit pools are pallid by comparison with those from shaded forest pools. These subjective comparisons were made with preserved specimens; detailed comparative data on living tadpoles are not available.

The relative length and depth of the tail are variable; in some individuals the greatest depth of the tail is about at mid-length of the tail, whereas in most specimens the tail is deepest at about one-third its length. The length of the tail relative to the total length is usually 58 to 64 per cent in tadpoles in stages 29 and 30 of development. In some individuals the tail is about 70 per cent of the total length. On the basis of the material examined, these variations in proportions do not show geographical trends. Probably the proportions are a reflection of crowding of the tadpoles in the pools where they are developing or possibly due to water currents or other environmental factors.

Stuart (1948:26) described and illustrated the tadpole of Smilisca baudini from Finca Chejel, Alta Verapaz, Guatemala. The description and figures agree with ours, except that the first lower tooth row does not have a sharp angle medially in Stuart's figure. He (1948:27) stated that color in tadpoles from different localities probably varies with soil color and turbidity of water. Maslin (1963:125) described and illustrated tadpoles of S. baudini from Pisté, Yucatán, México. These specimens are heavily pigmented like specimens that we have examined from the Yucatán Peninsula and from other places in the range of the species. Maslin stated that the anal tube is median in the specimens that he examined; we have not studied Maslin's specimens, but all tadpoles of Smilisca that we have examined have a dextral anal tube.

Newly metamorphosed young have snout-vent lengths of 12.0 to 15.5 mm. (average 13.4 in 23 specimens). The largest young are from La Libertad, El Petén, Guatemala; these have snout-vent lengths of 14.0 to 15.5 mm. (average 14.5 in five specimens). Young from 11 kilometers north of Vista Hermosa, Oaxaca, México, are the smallest and have snout-vent lengths of 12.0 to 12.5 mm. (average 12.3 in three specimens). Recently metamorphosed young usually are dull olive green above and white below; brown transverse bands are visible on the hind limbs. The labial markings characteristic of the adults are represented only by a creamy white suborbital spot, which is a good diagnostic mark for young of this species. In life the iris is pale gold.

Remarks: The considerable variation in color and the extensive geographic distribution of Smilisca baudini have resulted in the proposal of eight specific names for the frogs that we consider to represent one species. Duméril and Bibron (1841:564) proposed the name Hyla baudini for a specimen (MNHN 4798) from México. Smith and Taylor (1950:347) restricted the type locality to Córdoba, Veracruz, México, an area where the species occurs in abundance. Baird (1854:61) named Hyla vanvlieti from Brownsville, Texas, and (1859:35) labelled the figures of Hyla vanvlieti [= Hyla baudini] on plate 38 as Hyla vociferans, a nomen nudum. Cope (1862:359) named Hyla muricolor from Mirador, Veracruz, México, and (1865:194) used the name Smilisca daulinia for a skeleton that he employed as the basis for the cranial characters diagnostic of the genus Smilisca, as defined by him. Although we cannot be certain, Cope apparently inadvertently used daulinia for baudini, just as he used daudinii for baudini (1871:205). Brocchi (1877:125) named Hyla pansosana from Panzos, Alta Verapaz, Guatemala.

PLATE 1

Dorsal views of skulls of young Smilisca baudini: (A) recently metamorphosed young (KU 60026), snout-vent length 12.6 mm. ×23; (B) young (KU 85438), snout-vent length 32.1 mm. ×9.

PLATE 2

Skull of adult female Smilisca baudini (KU 68184): (A) Dorsal; (B) Ventral. ×4.5.

PLATE 3

Skull of adult female Smilisca baudini (KU 68184): (A) Lateral; (B) Dorsal view of left mandible; (C) Posterior. ×4.5.

PLATE 4

Palmar views of right hands of Smilisca: (A) S. baudini (KU 87177); (B) S. phaeota (KU 64276); (C) S. cyanosticta (KU 87199); (D) S. sordida (KU 91761); (E) S. puma (KU 91716), and (F) S. sila (KU 77408). ×3.

PLATE 5

Ventral aspect of right feet of Smilisca: (A) S. baudini (KU 87177); (B) S. phaeota (KU 64276); (C) S. cyanosticta (KU 87199); (D) S. sordida (KU 91761); (E) S. puma (KU 91716), and (F) S. sila (KU 77408). ×3.

PLATE 6

Living Smilisca: (A) S. baudini (UMMZ 115179) from 1.7 km. W Xicotencatl, Tamaulipas, México; (B) S. cyanosticta (UMMZ 118163) from Volcán San Martín, Veracruz, México; (C) S. phaeota (KU 64282) from Barranca del Río Sarapiquí, Heredia Prov., Costa Rica. All approx. nat. size.

PLATE 7

Living Smilisca: (A) S. puma (KU 65307) from 5.9 km. W. Puerto Viejo, Heredia Prov., Costa Rica; (B) S. sila (KU 77407) from Finca Palosanto, 6 km. WNW El Volcán, Chiriquí, Panamá; (C) S. sordida (KU 64257) from 20 km. WSW San Isidro el General, San José Prov., Costa Rica. All approx. nat. size.

PLATE 8

Fig. 1. Breeding site of Smilisca baudini, 4 km. WNW of Esparta, Puntarenas Prov., Costa Rica.

Fig. 2. Breeding site of Smilisca phaeota, Puerto Viejo, Heredia Prov., Costa Rica.

PLATE 9

Fig. 1. Breeding site of Smilisca puma, 7.5 km. W of Puerto Viejo, Heredia Prov., Costa Rica.

Fig. 2. Breeding site of Smilisca sordida, Río La Vieja, 30 km. E of Palmar Norte, Puntarenas Prov., Costa Rica.

PLATE 10

Audiospectrographs and sections of breeding calls of Smilisca: (A) S. baudini (KU Tape No. 74); (B) S. cyanosticta (KU Tape No. 373); (C) S. phaeota (KU Tape No. 79).

PLATE 11

Audiospectrographs and sections of breeding calls of Smilisca: (A) S. puma (KU Tape No. 382); (B) S. sila (KU Tape No. 385); (C) S. sordida (KU Tape No. 398).

PLATE 12

Lateral views of the heads of Smilisca: (A) S. baudini (KU 87177); (B) S. sordida (KU 91765); (C) S. phaeota (KU 64276); (D) S. puma (KU 91716); (E) S. cyanosticta (KU 87199); (F) S. sila (KU 77408). ×3.2.

Aside from the skeleton referred to as Smilisca daulinia by Cope (1865:194), we have examined each of the types of the species synonymized with S. baudini. All unquestionably are representatives of S. baudini.

Taylor (1942:306) named Hyla beltrani from Tapachula, Chiapas. This specimen (UIMNH 25046) is a small female (snout-vent length, 44 mm.) of S. baudini. Taylor (1954:630) named Hyla manisorum from Batán, Limón, Costa Rica. The type (KU 34927) is a large female (snout-vent length, 75.3 mm.) S. baudini. In this specimen and a male from Suretka, Costa Rica, the usual dorsal color pattern is absent, but the distinctive curved supraorbital processes, together with other structural features, show that the two specimens are S. baudini.

Hyla baudini dolomedes Barbour (1923:11), as shown by Dunn (1931a:413), was based on a specimen of Smilisca phaeota from Río Esnápe, Darién, Panamá.

Fig. 1. Map showing locality records for Smilisca baudini.

Distribution.—Smilisca baudini inhabits lowlands and foothills usually covered by xerophytic vegetation or savannas, but in the southern part of its range baudini inhabits tropical evergreen forest. The species ranges throughout the Pacific and Atlantic lowlands of México from southern Sonora and the Río Grande embayment of Texas southward to Costa Rica, where on the Pacific lowlands the range terminates at the southern limits of the arid tropical forest in the vicinity of Esparta; on the Caribbean lowlands the distribution seems to be discontinuous southward to Suretka (Fig. 1). Most localities where the species has been collected are at elevations of less than 1000 meters. Three localities are notably higher; calling males were found at small temporary ponds in pine-oak forest at Linda Vista, 2 kilometers northwest of Pueblo Nuevo Solistahuacán, Chiapas, elevation 1675 meters, and 10 kilometers northwest of Comitán, Chiapas, at an elevation of 1925 meters. Tadpoles and metamorphosing young were obtained from a pond in arid scrub forest, 17 kilometers northeast of Juchatengo, Oaxaca, elevation 1600 meters. Stuart (1954:46) recorded the species at elevations up to 1400 meters in the south-eastern highlands of Guatemala.

Specimens examined.—3006, as follows: United States: Texas: Cameron County, Brownsville, CNHM 5412-3, 6869, UMMZ 54036, USNM 3256.

Mexico: Campeche: Balchacaj, CNHM 102285, 102288, 102291, 102311, UIMNH 30709-22, 30726; Champotón, UMMZ 73172 (2), 73176, 73180; 16 km. E Champotón, UMMZ 73181; 5 km. S Champotón, KU 71369-75; 9 km. S Champotón, KU 71367-8; 10.5 km. S Champotón, KU 71365-6, 71722 (tadpoles), 71723 (yg.); 24 km. S Champotón, UMMZ 73177 (2); Chuina, KU 75101-3; Ciudad del Carmen, UIMNH 30703-8; Dzibalchén, KU 75413-31; Encarnación, CNHM 102282, 102289, 102294-5, 102300, 102306-8, 102312, 102314, 102316-7, 102319, 102322, UIMNH 30727-40, 30836-7; 1 km. W Escárcega, KU 71391-6; 6 km. W Escárcega, KU 71397-403; 7.5 km. W Escárcega, KU 71376-89; 14 km. W Escárcega, KU 71390; 13 km. W, 1 km. N Escárcega, KU 71404; 3 km. N Hopelchén, KU 75410-11; 2 km. NE Hopelchén, KU 75412; Matamoras, CNHM 36573; Pitál, UIMNH 30741; 1 km. SW Puerto Real, Isla del Carmen, KU 71345-64; San José Carpizo, UMMZ 99879; Tres Brazos, CNHM 102284, UIMNH 30723-5; Tuxpeña Camp, UMMZ 73239.

Chiapas: Acacoyagua, USNM 114487-92; 2 km. W Acacoyagua, USNM 114493-4; 5 km. E Arroyo Minas, UIMNH 9533-7; Berriozabal, UMMZ 119186 (7); Chiapa de Corzo, UMMZ 119185 (2); Cintalapa, UIMNH 50077; Colonia Soconusco, USNM 114495-9; 5 km. W Colonia Soconusco, UMMZ 87885 (7); Comitán, UMMZ 94438; 10 km. NW Comitán, KU 57185; El Suspiro, UMMZ 118819 (11); Escuintla, UMMZ 88271 (7), 88278, 88327, 109233; 6 km. NE Escuintla, UMMZ 87856 (26); 3 km. E Finca Juárez, UIMNH 9538; Finca Prussia, UMMZ 95167; Honduras, UMMZ 94434-7; La Grada, UMMZ 87862; 21 km. S La Trinitaria, UIMNH 9540-1; 14.4 km. SW Las Cruces, KU 64239-44; Palenque, UIMNH 49286, USNM 114473-84; 2 km. NW Pueblo Nuevo Solistahuacán, KU 57182-4, UMMZ 119948 (8), 121514; 1.3 km. N Puerto Madero, KU 57186-9; 4 km. N Puerto Madero, KU 57190-1; 8 km. N Puerto Madero, UMMZ 118379 (2); 12 km. N Puerto Madero, KU 57192; 17.6 km. N Puerto Madero, UMMZ 118378; Rancho Monserrata, UIMNH 9531-2, UMMZ 102266-7; Region Soconusco, UIMNH 33542-56; San Bartola, UIMNH 9519-30; San Gerónimo, UIMNH 30804; San Juanito, USNM 114485-6; San Ricardo, CNHM 102406; Solosuchiapa, KU 75432-3; Tapachula, CNHM 102208, 102219, 102239, 102405, UIMNH 25046, 30802-3; Tonolá, AMNH 531, CNHM 102232, 102416, UIMNH 30805-9, USNM 46760; Tuina, KU 41593 (skeleton); Tuxtla Gutierrez, CNHM 102231, 102248; 6 km. E Tuxtla Gutierrez, UIMNH 9539; 10 km. E Tuxtla Gutierrez, UMMZ 119949.

Chihuahua: 2.4 km. SW Toquina, KU 47226-7; Riito, KU 47228.

Coahuila: mountain near Saltillo, UIMNH 30833-4.

Colima: No specific locality, CNHM 1632; Colima, AMNH 510-11; Hacienda Albarradito, UMMZ 80029 (2); Hacienda del Colomo, AMNH 6208; Los Mezcales, UMMZ 80028; Manzanillo, AMNH 6207, 6209; Paso del Río, CNHM 102207, 102229-30, UIMNH 30819-21, UMMZ 110875 (3); Periquillo, UMMZ 80025 (3), 80026 (14); 1.6 km. SW Pueblo Juárez, UMMZ 115564; Queseria, CNHM 102204, 102216-7, 102224, UIMNH 30816-8, UMMZ 80023 (7), 80024 (7); Santiago, UMMZ 80027; 7.2 km. SW Tecolapa, UMMZ 115184.

Guerrero: Acahuizotla, UF 1338 (2), 1339-40, UMMZ 119182 (2), 119184; 3 km. S Acahuizotla, KU 87183-7; Acapulco, AMNH 55276, UMMZ 121879 (4), USNM 47909; 3 km. N Acapulco, UMMZ 110127; 8 km. NW Acapulco, UF 11203 (7); 27 km. NE Acapulco, UIMNH 26597-610; Agua del Obispo, CNHM 102214, 102290, 102293, 102310, 102413, KU 60413, 87180-2, UIMNH 30764-6; Atoyca, KU 87175-8; Buena Vista, CNHM 102279, 102304, 102313, 102315, UIMNH 30774; Caculutla, KU 87179; 20 km. S Chilpancingo, CNHM 102242, 102401, 102410-1, 102415; Colonia Buenas Aires, UMMZ 119189; El Limoncito, CNHM 102292, 102303, 102321, 102414; El Treinte, CNHM 102212, 102221, 102237, 102240-1, UIMNH 30783-5, USNM 114508-10; Laguna Coyuca, UMMZ 80960 (2); 3 km. N Mazatlán, UIMNH 30777-9; 9 km. S Mazatlán, CNHM 102209, 102215, 102234, 102246, UIMNH 30781-2; Mexcala, CNHM 102399, 102403, 102409, 106539-40, UIMNH 30775-6; Ocotito, KU 60414-23; 5.4 km. N Ocotito, UMMZ 119181 (4); 1.6 km. N Organos, UIMNH 30752-63; Palo Blanco, CNHM 102283, 102286, 102305, 102320, 102404, UIMNH 30767-70; Pie de la Cuesta, AMNH 55275, 59202-5; Puerto Marquéz, AMNH 59200-1 (13); 5.6 km. S San Andreas de la Cruz, KU 87173-4; San Vincente, KU 87172; Zaculapán, UMMZ 119183.

Hidalgo: Below Tianguistengo, CNHM 102318.

Jalisco: Atenqueque, KU 91435-6; 5 km. NE Autlán, UIMNH 30810; 5 km. E Barro de Navidad, UMMZ 110900; Charco Hondo, UMMZ 95247; Puerto Vallarta, UIMNH 41346; between La Huerta and Tecomates, KU 91437; 3 km. SE La Resolana, KU 27619, 27620 (skeleton); 11 km. S, 1.6 km. E Yahualica, KU 29039; Zapotilitic, CNHM 102238.

Michoacán: Aguililla, UMMZ 119179 (5); Apatzingán, CNHM 38766-90, KU 69101 (skeleton); 7 km. E Apatzingán, UMMZ 112843; 11 km. E Apatzingán, UMMZ 112841 (3); 27 km. S Apatzingán, KU 37621-3; 1.6 km. N Arteaga, UMMZ 119180; Charapendo, UMMZ 112840; Coahuayana, UMMZ 104458; El Sabino, CNHM 102205-6, 102210-1, 102220, 102228, 102233, UIMNH 30822-3; La Placita, UMMZ 104456; La Playa, UMMZ 105163; 30 km. E Nueva Italia, UMMZ 120255 (2); 4 km. S Nueva Italia, UMMZ 112842; Ostula, UMMZ 104457 (4); Salitre de Estopilas, UMMZ 104459; San José de la Montaña, UMMZ 104461 (2); 11 km. S Tumbiscatio, KU 37626; 12 km. S Tzitzio, UMMZ 119178.

Morelos: 3.5 km. W Cuautlixco, KU 87188-90; 1 km. NE Puente de Ixtla, KU 60393-4; 20 km. S Puente de Ixtla, CNHM 102400, UIMNH 30832; Tequesquitengo, AMNH 52036-9.

Nayarit: 3 km. S Acaponeta, UMMZ 123030 (4); 56 km. S Esquinapa (Sinaloa), KU 73909; Jesús María, AMNH 58239; San Blas, KU 28087, 37624, 62360-2, USNM 51408; 8.6 km. E San Blas, UMMZ 115185; Tepic, UIMNH 30812-5; 4 km. E Tuxpan, KU 67786; 11 km. SE Tuxpan, UIMNH 7329-31, 7335-59.

Nuevo León: Galeana, CNHM 34389; Salto de Cola de Caballo, CNHM 30628-31, 30632 (40), 30633-7, 34454-67.

Oaxaca: 11 km. S Candelaria, UIMNH 9515-8; Cerro San Pedro, 24 km. SW Tehuantepec, UMMZ 82156; Chachalapa, KU 38199; 8 km. S Chiltepec, KU 87191; 12 km. S Chivela, UMMZ 115182; Coyul, USNM 114512; Garza Mora, UIMNH 40967-8; Juchatengo, KU 87193; 17 km. NE Juchatengo, KU 87645 (tadpoles), 87646 (young); Juchitán, USNM 70400; Lagartero, UIMNH 9514; Matías Romero, AMNH 52143-5; 25 km. N Matías Romero, KU 33822-8; 7 km. S Matías Romero, UIMNH 42703; Mirador, AMNH 6277, 13832-9, 13842-55; Mira León, 1.6 km. N Huatulco, UIMNH 9503-4; Mixtequillo, AMNH 13924; Pochutla, KU 57167-81, UIMNH 9505-13; Quiengola, AMNH 51817, 52146; Río del Corte, UIMNH 48677; Río Mono Blanco, UIMNH 36831; Río Sarabia, 5 km. N Sarabia, UMMZ 115180 (4); 2.5 km. N Salina Cruz, KU 57165-6; San Antonio, UIMNH 37286; 5 km. NNW San Gabriel Mixtepec, KU 87192; San Pedro del Istmo, UIMNH 37197; Santo Domingo, USNM 47120-2; 3.7 km. N Sarabia, UMMZ 115181 (3); Tapanatepec, KU 37793 (skeleton), 37794, UIMNH 9542, UMMZ 115183; between Tapanatepec and Zanatepec, UIMNH 42704-25; Tecuane, UMMZ 82163 (3); Tehuantepec, AMNH 52625, 52639, 53470, UMMZ 82157-8, 82159 (9), 82160 (4), 82161 (8), 82162 (12), 112844-5, 118703, USNM 10016, 30171-4, 30188; 4.5 km. W Tehuantepec, KU 59801-12 (skeletons), 69102-3 (skeletons); 10 km. S Tehuantepec, KU 57163-4; Temazcal, USC 8243 (3); 3 km. S Tolocita, KU 39666-9; Tolosa, AMNH 53605; Tuxtepec, UMMZ 122098 (2); 2 km. S Valle Nacional, KU 87194-5; 11 km. N Vista Hermosa, KU 87196, 87639-41 (tadpoles), 87642-3 (young), 87644 (tadpoles); Yetla, KU 87197.

Puebla: 16 km. SW Mecatepec (Veracruz), UIMNH 3657-8; San Diego, AMNH 57714, USNM 114511; Vegas de Suchil, AMNH 57712; Villa Juárez, UF 11205.

Quintana Roo: Cóba, CNHM 26937; Esmeralda, UMMZ 113551; 4 km. NNE Felipe Carrillo Puerto, KU 71417-8; Pueblo Nuevo X-Can, KU 71405; 10 km. ENE Pueblo Nuevo X-Can, KU 71406; 4 km. WSW Puerto Juárez, KU 71407-11, 71721 (tadpoles); 12 km. W Puerto Juárez, KU 71412-6; San Miguel, Isla de Cozumel, UMMZ 78542 (6), 78543 (10), 78544 (2); 3.5 km. N San Miguel, Isla de Cozumel, KU 71419-22; 10 km. E San Miguel, Isla de Cozumel, UMMZ 78541; Telantunich, CNHM 26950.

San Luis Potosí: Ciudad Valles, AMNH 57776-81 (12), CNHM 37193, 102297, KU 23705; 21 km. N Ciudad Valles, UMMZ 118377; 6 km. E Ciudad Valles, UF 3524; 24 km. E Ciudad Valles, UF 7340 (2); 5 km. S Ciudad Valles, UIMNH 30751; 16 km. S Ciudad Valles, AMNH 52953; 30 km. S Ciudad Valles, CNHM 102394, 102402, 102412, UIMNH 30749-50; 63 km. S Ciudad Valles, UIMNH 19247-58; Pujal, UMMZ 99872 (2); Río Axtla, near Axtla, AMNH 53211-5, 59516, KU 23706; Tamazunchale, AMNH 52675, CNHM 39621-2, 102226, 102281, UF 7615 (2), UIMNH 26596, UMMZ 99506 (9), 118701 (2), USNM 114468; 17 km. N Tamazunchale, UIMNH 3659; 2.4 km. S Tamazunchale, AMNH 57743; 17 km. E Tamuin, UF 11202 (2); Xilitla, UIMNH 19259-60.

Sinaloa: 8 km. N. Carrizalejo, KU 78133; 4 km. NE Concordia, KU 73914; 5 km. SW Concordia, KU 75438-9; 6 km. E Cosalá, KU 73910; Costa Rica, 16 km. S. Culiacán, UIMNH 34887-9; 51 km. SSE Culiacán, KU 37792; El Dorado, KU 60392; 1.6 km. NE El Fuerte, CNHM 71468; Isla Palmito del Verde, middle, KU 73916-7; 21 km. NNE Los Mochis, UIMNH 40536-7; Matatán, KU 73913; 7.3 km. SW Matatán, KU 78464, 78466-70; Mazatlán, AMNH 12562, UMMZ 115197 (3); 57 km. N Mazatlán, UIMNH 38364; Plomosas, USNM 47439-40; Presidio, UIMNH 30811, USNM 14082; Rosario, KU 73911-2; 5 km. E Rosario, UIMNH 7360-76; 8 km. SSE Rosario, KU 37625; 5 km. SW San Ignacio, KU 78465; 1.6 km. ENE San Lorenzo, KU 47917-24; Teacapán, Isla Palmito del Verde, KU 73915; 9.6 km. NNW Teacapán, KU 91410; Villa Unión, KU 78471; 9 km. NE Villa Unión, KU 75434-7; 1 km. W Villa Unión, AMNH 59284.

Sonora: Guiracoba, AMNH 51225-38 (25).

Tabasco: 4 km. NE Comalcalco, AMNH 60313; Teapa, UMMZ 119943; 5 km. N Teapa, UMMZ 119940, 119944, 122997 (2); 10 km. N Teapa, UMMZ 119187, 119188 (2); 13 km. N Teapa, UMMZ 119941 (2), 119945 (3), 120254 (2); 21 km. N Teapa, UMMZ 119942, 119947; 29 km. N Teapa, UMMZ 119946 (11); Tenosique, USNM 114505-7.

Tamaulipas: Acuña, UMMZ 99864; 5 km. S Acuña, 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; Gómez Farías, UMMZ 110837-8; 8 km. NE Gómez Farías, UMMZ 102265, 102916 (4), 102917, 104110 (5), 105493, 110836 (2), 111274-7; 8 km. NW Gómez Farías, UMMZ 101178 (7), 101179 (3), 101362-3, 101364 (2), 108799 (2), 110129, 111278, 111280; 8 km. W Gómez Farías, UMMZ 102859 (2); 16 km. W Gonzales, KU 37795-6; Jiménez, KU 60412; La Clementina, 6 km. W Forlan, USNM 106244; Limón, 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; Villagrán, 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 Amatitlán, UMMZ 115195; Barranca Metlac, UIMNH 38365; Boca del Río, UIMNH 26619-30, UMMZ 74954 (9); 16 km. S Boca del Río, UIMNH 26631; between Boca del Río and Anton Lizardo, UIMNH 42701; Canadá, CNHM 102397; Catemaco, UMMZ 118702 (4); Ciudad Alemán, UMMZ 119608 (3); Córdoba, CNHM 38665-7, USNM 30410-3; 5.2 km. ESE Córdoba, KU 71423-35, 89924 (skeleton); 7 km. ESE Córdoba, 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 Ríos, 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 Fortín, UF 11201; 4 km. SW Huatusco, UMMZ 115177; 1 km. SW Huatusco, UMMZ 123119; 10 km. SE Hueyapan, UMMZ 115190; 20 km. S Jesús Carranza, KU 23844, 23846, 27399; 38 km. SE Jesús Carranza, KU 23417; Laguna Catemaco, UMMZ 119932 (62); 1.6 km. N La Laja, UIMNH 3651; La Oaxaqueña, AMNH 43930-1; 17 km. E Martínez de la Torre, UIMNH 36630-2; 6.2 km. W Martínez de la Torre, UIMNH 3652-4; Minatitlán, 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; Pérez, CNHM 1686 (5); 20 km. N Piedras Negras, Río Blanco, KU 23708; Plan del Río, 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); Río de las Playas, USNM 118635-6; Río Seco, UMMZ 88801 (9); Rodriguez Clara, CNHM 102225; San Andrés Tuxtla, CNHM 102213, 102222, 102227, 102247, UIMNH 30789-91; 10 km. NW San Andrés Tuxtla, UMMZ 119935; 13.4 km. NW San Andrés Tuxtla, UMMZ 119939 (2); 19.8 km. NW San Andrés Tuxtla, UMMZ 119938; 27.2 km. NW San Andrés Tuxtla, UMMZ 119936 (6); 48 km. NW San Andrés Tuxtla, UMMZ 119937; 4 km. W San Andrés Tuxtla, UMMZ 115187; 37.4 km. S San Andrés 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 Tehuatlán, UIMNH 3660-3; 5 km. S Tehuatlán, 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.

Yucatán: No specific locality, CNHM 548, 49067, USNM 32298; Chichén-Itzá, 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 Chichén-Itzá, KU 71438-9; 12 km. E Chichén-Itzá, KU 71440; Mérida, CNHM 40659-66, UIMNH 30747-8, UMMZ 73182; 6 km. S Mérida, 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); Chamá, MCZ 15792-3, UMMZ 90895 (7), 90896 (5), 90897 (29), 90898 (12), 90899; Chinajá, KU 55939-41, 57193-8, 60018-20 (tadpoles), 60021 (eggs), 60022 (tadpoles); Cobán, 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 Volcán, 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 Gerónimo, UMMZ 84076 (16).

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

El Petén: 20 km. NNW Chinajá (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; Poptún, UMMZ 120475; Poza de la Jicotea, USNM 114672; Ramate-Yaxha trail, UMMZ 75321; Río de la Pasión between Sayaxché and Subín, KU 57151; Río San Román, 16 km. NNW Chinajá (Alta Verapaz), KU 55942-6; Sacluc, USNM 25131; Sayaxché, KU 57144-5; Tikal, UMMZ 117983 (7), 117984 (5), 117993 (5), 120474 (5); Toocog, KU 57241-2, 60023 (young), 60025 (young); Uaxactún, UMMZ 70401-3; Yaxha, UMMZ 75322; 19 km. E Yaxha, UMMZ 75330 (4).

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

Escuintla: Río 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).

Izabál: 2 km. SW Puerto Matías de Gálvez, KU 60027-8 (tadpoles); Quiriguá, 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. Morazán: 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 Roatán, 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). Colón: Bambú, UF 320; Belfate, AMNH 45692-5; Patuca, USNM 20261. Comayagua: La Misión, 3.5 leagues N Siguatepeque, MCZ 26424-5. Copán: Copán, UMMZ 83026 (2). Cortés: Cofradía, AMNH 45345-6; Hacienda Santa Ana, CNHM 4724-31; Lago de Yojoa, MCZ 26410-1; Río Lindo, AMNH 54972. El Paraiso: El Volcán, MCZ 26436. Francisco Morazán: Tegucigalapa, BYU 18301-4, 18837-41, MCZ 26395-7, USNM 60499. Gracias A Dios: Río Segovia, MCZ 24543. Santa Barbara: Santa Barbara, USNM 128062-5.

Nicaragua: Chinandega: 4 km. N, 2 km. W Chichigalpa, KU 85385; Chinandega, MCZ 2632; Río Tama, USNM 40022; San Antonio, KU 84944-9 (skeletons), 85386-403. Chontales: 1 km. NE Acoyapa, KU 64234. Estelí: Finca Daraili, 5 km. N, 15 km. E Condega, KU 85404-8; Finca Venecia, 7 km. N, 16 km. E Condega, KU 85409. León: 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. Río San Juan: Greytown, USNM 19585-6, 19767-8. Rivas: Javillo, UMMZ 123001; Moyogalpa, Isla Ometepe, KU 85428-37, 87706 (tadpoles); Peñas Blancas, KU 85417; Río Javillo, 3 km. N, 4 km. W Sapoá, 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, Río Huaspuc, AMNH 58428-31; 11 km. NW Rama, Río Siquia, UMMZ 79708, 79709 (5), 79710 (2); Río Escondido, USNM 19766, 20701; Río Siquia at Río 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 Cañas, 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; Río Piedra, 1.6 km. W Bagaces, USC 7027; Río Bebedero, 5 km. S Bebedero, KU 64158; 5 km. NE Tilarán, KU 36782-6. Heredia: 13 km. SW Puerto Viejo, KU 64142-6. Limón: Batán, 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 Petén, 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 Petén, 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 Chinajá, 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 Chinajá, 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 Chinajá (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, México, 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 Volcán San Martin, Veracruz, México, 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 Chinajá, 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 Volcán San Martín, Veracruz, México, 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 Chinajá, 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, México, on June 23-28, 1964; in a quiet pool in a stream 8 kilometers south of Yetla, Oaxaca, México, in July, 1963 (Dale L. Hoyt, personal communication); in and near springs flowing into a stream at Dos Amates, Veracruz, México, on August 4, 1959 (Douglas Robinson, personal communication); and in a water-filled depression in a log 10 kilometers west-northwest of Chinajá, 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 Volcán San Martín, Veracruz, México, 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 Chinajá, 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, México) 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, México): "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 México and northern Central America from northern Oaxaca and southern Veracruz through northern Chiapas in México and into El Petén and northern Alta Verapaz in Guatemala (Fig. 2). Apparently the range is discontinuous, for in southern México the species is found in cloud forest at elevations of 830 to 900 meters on the northern slopes of the Sierra de Juárez. 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 Petén, 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 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 for Smilisca cyanosticta (triangles) and Smilisca phaeota (circles).

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. 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 labialis Peters, 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 dolomedes Barbour, 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 distinguishing S. phaeota from 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 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 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 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, 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 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, 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 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 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, 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 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) named Hyla labialis from the "region of Bogotá, 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 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 combination

Hyla puma Cope, 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 wellmanorum Taylor, 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 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 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, 0.06-0.35 seconds; of secondary notes, 0.10 to 0.47 seconds), repeated at intervals of 5 to 55 seconds. The primary notes have 187 to 240 pulses per second and major frequencies of about 740 to 1870 cycles per second (Pl. 11A).

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

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

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

Remarks.—The identity of Cope's Hyla puma has not been known. The name has appeared in various compilations, but no workers have referred any of their specimens to that species. Examination of the holotype (USNM 13735), an adult female, revealed the presence of the following combination of characters: snout-vent length 45.8 mm., snout blunt above and rounded laterally, nostrils close to tip of snout, lips thin and flaring, a vestige of a web on the hands, feet about one-half webbed, tarsal fold weak and extending about two-thirds length of tarsus, dorsal markings consisting of a faded dark interorbital bar and a pair of faded longitudinal brown marks connected by a transverse band in the scapular region. The type agrees well with specimens of Smilisca wellmanorum (Taylor, 1952); the vestigial webbing on the hands and the dorsal coloration are especially significant. Consequently, we consider Hyla wellmanorum Taylor, 1952, to be a synonym of Hyla puma Cope, 1885. Cope gave only "Nicaragua" as the locality for Hyla puma. The specimen was part of a collection received at the United States National Museum from Lt. J. F. Moser. Among the species in the collection are Dentrobates pumilio, Phyllomedusa helenae, Corythophanes cristatus, Pliocercus dimidatus, Tretanorhinus nigroluteus, and others characteristically found on the Caribbean lowlands of Central America. Thus, it seems reasonable to assume that the type specimen of Hyla puma came from the Caribbean lowlands. Though no other Nicaraguan specimens have been found by us, numerous specimens are known from the Caribbean lowlands of Costa Rica.

Cochran (1961:58), in her catalogue of type specimens in the United States National Museum, listed Hyla puma Cope, 1885, as a synonym of Hyla molitor Schmidt, 1857. She made no qualifying statements. Schmidt (1858:246), in his descriptions of the species in the year following his publication of the names and Latin diagnoses, stated: "Dorsum uniformly gray, more intensive on back, fading away laterally and on extremities; in every-day-life this blue would be called Mueller's Blau. A delicately dotted black line runs on the canthus rostralis from the opening of the nose to the corner of the eye. In the armpits, on the flanks and the thighs two of our three specimens have black marblings." [Free translation] Certainly on the basis of coloration Hyla puma is distinctly different from Hyla molitor.

Distribution.—This species lives in the wet, forested regions of the Caribbean lowlands of Costa Rica and presumably southern Nicaragua (Fig. 3). All specimens are from low elevations; the highest known elevation for the occurrence of this frog is 285 meters at Laguna Bonilla.

Fig. 3. Map showing locality records for Smilisca puma (triangles) and Smilisca 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 Puerto Viejo, 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 sila new 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 distinguishing S. sila from 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 of Smilisca sila consists 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 in Smilisca sordida. In one specimen (KU 80467) the blotches are fused and form two wide irregular longitudinal stripes, as in Smilisca 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 from Costa 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 sila inhabits 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; musculature extending 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 with Smilisca sordida; most authors have referred both species to Hyla (Smilisca) gabbi. Examination of the types of Hyla sordida, gabbi, salvini, and nigripes revealed 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. Possibly Hyla molitor Schmidt (1857) is based on the species that we have named S. 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 sila ranges 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 the Caribbean lowlands and in the valleys of the northward draining rivers of Colombia (Fig. 3).

Specimens examined, 234, as follows: Costa Rica: Puntarenas: 6 km. E Golfito, KU 91717; Quebrada Boruca, 22 km. E Palmar Norte, KU 64265-6; Río Zapote, 7 km. E Palmar Norte, USC 7100 (2). San José: San Isidro el General, KU 28200; 14 km. NW San Isidro el General, USC 7098 (2); 15 km. WSW San Isidro el General, USC 7097.

Panama: Canal Zone: Barro Colorado Island, AMNH 62320-3, CNHM 13324, 13326-8, 13330, 13338, 13359, 13423-5, KU 80460-6, 80619 (young), 80625 (skeleton), UMMZ 63542-6, USC 7051. Chiriquí: Boquete, AMNH 69815, UMMZ 58441-5; El Volcán, KU 77413, 91828-31 (skeletons), 91852-74, 91832 (eggs), 91833 (tadpoles); 6 km. S El Volcán, CNHM 60442; 16 km. NNW El Volcán, KU 91879-90; Finca Palosanto, 6 km. WNW El Volcán, [Pg 323] KU 77406-12, 77692 (skeleton), 91875-7, 92330-1; Río Colorado, 17 km. NNW El Volcán, KU 91878, 92332; Valle Hornito, 19 km. NE Gualaca, KU 92333-45. Coclé: El Valle, AMNH 55440-5 (13), 59607-14, CNHM 48140, 60349-2, 60387-92, 60401-4, 60443, 67842-5, KU 91834 (young), 91902-4, TNHC 23751-2, USNM 140653. Colón: Río Candelaria, AMNH 53708-15, CNHM 67826-36. Darién: Camp Creek, Camp Townsend, AMNH 40756-7, 40936-9, 40992; Río Chico, AMNH 39784, 40986-7; Río Pita, CNHM 67823-5; Tacarcuna, USNM 141796-802; Three Falls Creek, AMNH 41684, 51788. Los Santos: Cerro Hoya, USNM 148213-4; Lajamina, Río Puria, KU 67915. Panamá: Altos de Pacora, KU 91894; Cerro Jefe, KU 91895-6; Cerro La Campana, CNHM 67846, KU 91897-900, USNM 139689; Finca La Sumbadora, KU 80467-81, 80620 (tadpoles), 91910 (eggs), 91911-2 (tadpoles), 91913 (young), 91908-9 (skeletons); Río Calobra, USNM 53722, Río Pacora, 9 km. NNE Pacora, KU 91901. Veraguas: Cerro Carbunco, USNM 129066; Cerro Tute, CNHM 67837-41; Isla Cebaco, Río Platanal, KU 91891-3.

Colombia: Antioquia: Urabá, Villa Arteaga, CNHM 63893 (Goin). Atlantico: Sabanalarga, Río Causa, AMNH 14506.

Smilisca sordida (Peters), new combination

Hyla sordida Peters, Monatsb. Konigl. Akad. Wissen. Berlin., p. 460, 1863 [Syntypes.—ZMB 3141 (two specimens) from "Veragua," Panamá; J. von Warszewicz collector]. Brocchi, Mission scientifique au Mexique ..., pt. 3, sec. 2, Études sur les batrachiens, p. 42, 1881. Boulenger, Catalogue Batrachia Salientia in British Museum, p. 393, Feb. 1, 1882. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p. 273, Sept. 1901. Nieden, Das Tierreich, Amphibia, Anura, I, p. 258, June, 1923.

Hyla gabbi Cope, Jour. Acad. Nat. Sci. Philadelphia, new ser., 8, pt. 2:103, 1876 [Syntypes.—USNM 30658-9 from near Sipurio, Limón, Costa Rica; William M. Gabb collector]. Brocchi, Mission scientifique au Mexique ..., pt. 3, sec. 2, Études sur les batrachiens, p. 37, 1881. Boulenger, Catalogue Batrachia Salientia in British Museum, p. 372, Feb. 1, 1882. Cope, Bull. U. S. Natl. Mus., 32:32, 1887. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p. 274, Sept. 1901. Werner, Abhand. Konigl. Akad. Wissen. München., 22:351, 1903. Nieden, Das Tierreich, Amphibia, Anura I, p. 252, June, 1923. Taylor, Univ. Kansas Sci. Bull., 35(1):840, July 1, 1952. Cochran, Bull. U. S. Natl. Mus., 220:54, 1961.

Hyla nigripes Cope, Jour. Acad. Nat. Sci. Philadelphia, new ser., 8, pt. 2:104, 1876 [Syntypes.—USNM 30685-6, from Pico Blanco, Costa Rica; William M. Gabb collector]. Brocchi, Mission scientifique au Mexique ..., pt. 3, sec. 2, Études sur les Batrachiens, p. 38, 1881. Boulenger, Catalogue Batrachia Salientia in British Museum, p. 394, Feb. 1, 1882. Cope, Bull. U. S. Natl. Mus., 32:32, 1887. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p. 278, Sept., 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 253, June, 1923. James, Copeia, 3:147, Sept. 30, 1944. Taylor, Univ. Kansas Sci. Bull, 35(1):853, July 1, 1952. Cochran, Bull. U. S. Natl. Mus., 220:56, 1961.

Hyla salvini Boulenger, Catalogue Batrachia Salientia in British Museum, p. 372, Feb. 1, 1882 [Syntypes.—BMNH 1947.2.24.13-14 from Cartago, Costa Rica; Osbert Salvin collector]. Günther, Biologia Centrali-Americana: Reptilia and Batrachia, pl. 71, Fig. B., Sept., 1901. Werner, Abhand. Zool.-Bot. Gesell. Wien, 46:8, Sept. 30, 1896.

Smilisca gabbi, Starrett, Copeia, 4:303, Dec. 30, 1960.

Diagnosis.—Size moderate ([M] 45 mm., [F] 64 mm.); skull slightly wider than long, having large and elongate frontoparietal fontanelle; supraorbital flanges absent; squamosal small, not contacting maxillary; bony section of ethmoid terminating just anterior to anterior edge of orbit; tarsal fold weak, full length of tarsus; inner metatarsal tubercle long, low, flat, elliptical; lips thin and flaring; fingers one-half webbed; toes four-fifths webbed; diameter of tympanum about one-half that of eye; no white labial stripe; dorsal dark markings irregular, sometimes forming broad transverse bars; pale flecks on flanks and usually on posterior surfaces of thighs; vocal sacs in breeding males white. (Foregoing combination of characters distinguishing S. sordida from any other species in genus.)