University of Kansas PublicationsMuseum of Natural HistoryVolume 17, No. 7, pp. 281-375, pls. 1-12, 17 figs.July 14, 1966Neotropical Hylid Frogs, Genus SmiliscaBYWILLIAM E. DUELLMAN AND LINDA TRUEBUniversity of Kansas Lawrence1966University of Kansas Publications, Museum of Natural HistoryEditors: E. Raymond Hall, Chairman, Henry S. Fitch, Frank B. CrossVolume 17, No. 7, pp. 281-375, pls. 1-12, 17 figs.Published July 14, 1966University of KansasLawrence, KansasPRINTED BYROBERT R. (BOB) SANDERS, STATE PRINTERTOPEKA, KANSAS1966Look for the Union Label31-3430Neotropical Hylid Frogs, Genus SmiliscaBYWILLIAM E. DUELLMAN AND LINDA TRUEBCONTENTSPAGEIntroduction285Acknowledgments286Materials and Methods287Genus Smilisca Cope, 1865287Key to Adults288Key to Tadpoles289Accounts of Species289Smilisca baudini(Duméril and Bibron)289Smilisca cyanosticta(Smith)303Smilisca phaeota(Cope)308Smilisca puma(Cope)314Smilisca silaNew species318Smilisca sordida(Peters)323Analysis of Morphological Characters330Osteology330Descriptive Osteology of Smilisca baudini331Developmental Cranial Osteology of Smilisca baudini333Comparative Osteology336Musculature341Skin342Structure342Comparative Biochemistry of Proteins343External Morphological Characters343Size and Proportions343Shape of Snout344Hands and Feet344Ontogenetic Changes344Coloration344Metachrosis345Chromosomes345Natural History345Breeding345Time of Breeding345Breeding Sites346Breeding Behavior346Breeding Call351Eggs356Tadpoles357General Structure357Comparison of Species357Growth and Development361Behavior365Phylogenetic Relationships366Interspecific Relationships366Evolutionary History369Summary and Conclusions371Literature Cited372INTRODUCTIONThe 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.Hylaand 10 other genera inhabit Central America; four of those 10 genera (Gastrotheca,Hemiphractus,Phrynohyas, andPhyllomedusa) are widely distributed in South America. The other six genera are either restricted to Central America or have their greatest differentiation there.PlectrohylaandPtychohylainhabit streams in the highlands of southern Mexico and northern Central America;DiaglenaandTriprionare casque-headed inhabitants of arid regions in México and northern Central America.Anothecais a tree-hole breeder in cloud forests in Middle America. The genusSmiliscais 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 generaDiaglena,Triprion,Tetraprion,Osteocephalus,Trachycephalus,Aparasphenodon,Corythomantis,Hemiphractus,Pternohyla, andAnothecahave been recognized as distinct from one another and from the genusHylaon the basis of various modifications of dermal bones of the cranium.Phyllomedusais recognized on the basis of a vertical pupil and opposable thumb;Plectrohylais characterized by the presence of a bony prepollex and the absence of a quadratojugal.Gastrothecais 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 distinguishPhrynohyasfromHyla. He (1963a) cited the ventrolateral glands in breeding males as diagnostic ofPtychohyla. Some species groups within the vaguely defined genusHylahave equally distinctive characters. TheHyla septentrionalisgroup is characterized by a casque-head, not much different from that in the genusOsteocephalus(Trueb, MS). Males in theHyla maximagroup have a protruding bony prepollex like that characteristically found inPlectrohyla.Ontogenetic development, osteology, breeding call, behavior, and ecology are important in the recognition of species. By utilizingthe combination of many morphological and biological factors, the genusSmiliscacan be defined reasonably well as a natural, phyletic assemblage of species. Because the wealth of data pertaining to the morphology and biology ofSmiliscais lacking for most other tree frogs in Middle America it is not possible at present to compareSmiliscawith related groups in more than a general way.Smiliscais 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 genusSmilisca, 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 genusSmiliscato 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.AcknowledgmentsExamination 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 MethodsIn 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 andcleanedand stained specimens of all species, plus serial sections of the skull ofSmilisca 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:AMNH—American Museum of Natural HistoryBMNH—British Museum (Natural History)BYU—Brigham Young UniversityCNHM—Chicago Natural History MuseumKU—University of Kansas Museum of Natural HistoryMCZ—Museum of Comparative ZoologyMNHN—Museu National d'Histoire Naturelle, ParisUF—University of Florida CollectionsUIMNH—University of Illinois Museum of Natural HistoryUMMZ—University of Michigan Museum of ZoologyUSC—University of Southern CaliforniaUSNM—United States National MuseumTNHC—Texas Natural History Collection, University of TexasZMB—Zoologisches Museum BerlinGenus SmiliscaCope, 1865SmiliscaCope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct., 1865 [Type speciesSmilisca dauliniaCope, 1865 =Hyla baudiniDuméril 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 mandibulaeconsisting of two parts, one arising from dorsal fascia and other from posterior arm of squamosal, (6) dividedM. 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 genusSmiliscacontains six recognizable species. An alphabetical list of the specific and subspecific namesthat we consider to be applicable to species ofSmiliscarecognized herein is given below.Names proposedValid namesHyla baudiniDuméril and Bibron, 1841=S. baudiniHyla baudini dolomedesBarbour, 1923=S. phaeotaHyla beltraniTaylor, 1942=S. baudiniHyla gabbiCope, 1876=S. sordidaHyla labialisPeters, 1863=S. phaeotaHyla manisorumTaylor, 1954=S. baudiniHyla muricolorCope, 1862=S. baudiniHyla nigripesCope, 1876=S. sordidaHyla pansosanaBrocchi, 1877=S. baudiniHyla phaeotaCope, 1862=S. phaeotaHyla phaeota cyanostictaSmith, 1953=S. cyanostictaHyla pumaCope, 1885=S. pumaHyla salviniBoulenger, 1882=S. sordidaHyla sordidaPeters, 1863=S. sordidaHyla vanvlietiiBaird, 1854=S. baudiniHyla vociferansBaird, 1859=S. baudiniHyla wellmanorumTaylor, 1952=S. pumaDistribution 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 Adults1.Larger frogs ([M] to 76 mm., [F] to 90 mm.) having broad flat heads and a dark brown or black postorbital mark encompassing tympanum2Smaller frogs ([M] to 45 mm., [F] to 64 mm.) having narrower heads and lacking a dark brown or black postorbital mark encompassing tympanum42.Lips barred; flanks cream-colored with bold brown or black mottling in groin; posterior surfaces of thighs brown with cream-colored flecksS. baudini, p. 289Lips not barred; narrow white labial stripe present; flanks not cream-colored with bold brown or black mottling in groin; posterior surfaces of thighs variable33.Flanks and anterior and posterior surfaces of thighs dark brown with large pale blue spots on flanks and small blue spots on thighsS. cyanosticta, p. 303Flanks cream-colored with fine black venation; posterior surfaces of thighs pale brown with or without darker flecks or small cream-colored spotsS. phaeota, p. 3084.Fingers having only vestige of web; diameter of tympanum two-thirds that of eye; dorsum pale yellowish tan with pair of broad dark brown stripesS. puma, p. 314Fingers about one-half webbed; diameter of tympanum about one-half that of eye; dorsum variously marked with spots or blotches55.Snout short, truncate; vocal sacs in breeding males dark gray or brown; blue spots on flanks and posterior surfaces of thighsS. sila, p. 318Snout long, sloping, rounded; vocal sacs in breeding males white; cream-colored or pale blue flecks on flanks and posterior surfaces of thighsS. sordida, p. 323Key to Tadpoles1.Pond tadpoles; tail about half again as long as body; mouth anteroventral2Stream tadpoles; tail about twice as long as body; mouth ventral52.Labial papillae in two rows3Labial papillae in one row43.First upper tooth row strongly arched medially; third lower tooth row much shorter than other rows; dorsal fin deepest at about two-thirds length of tail; tail cream-colored with dense gray reticulationsS. puma, p. 314First upper tooth row not arched medially; third lower tooth row nearly as long as others; dorsal fin deepest at about one-third length of tail; tail tan with brown flecks and blotchesS. baudini, p. 2894.Dorsal fin extending onto bodyS. phaeota, p. 308Dorsal fin not extending onto bodyS. cyanosticta, p. 3035.Mouth completely bordered by two rows of papillae; inner margin of upper beak not forming continuous arch with lateral processes; red or reddish brown markings on tailS. sordida, p. 323Median part of upper lip bare; rest of mouth bordered by one row of papillae; inner margin of upper beak forming continuous arch with lateral processes; dark brown markings on tailS. sila, p. 318ACCOUNTS OF SPECIESSmilisca baudini(Duméril and Bibron)Hyla baudiniDuméril 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 vanvlietiiBaird, 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 vociferansBaird, 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 muricolorCope, 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 dauliniaCope, 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 forbaudini], 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 pansosanaBrocchi, 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 beltraniTaylor, 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 manisorumTaylor, 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 distinguishingS. baudinifrom 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.Table 1.—Geographic Variation in Size and Proportions in Males of Smilisca baudini. (Means in Parentheses Below Observed Ranges; Data Basedon10 Specimens From Each Locality.)LocalitySnout-ventlengthTibia length/snout-ventTympanum/eyeSouthern Sinaloa62.3-75.943.2-46.784.2-94.4(68.6)(44.9)(87.8Ocotito, Guerrero55.6-64.046.1-51.266.7-82.8(58.7)(47.8)(74.6)Pochutla, Oaxaca56.1-65.144.7-49.473.0-84.2(60.2)(47.5)(77.4)San Salvador, El Salvador57.0-68.042.1-46.174.6-83.3(62.1)(44.9)(77.6)Managua, Nicaragua52.9-63.645.6-49.473.7-89.7(57.3)(47.5)(79.4)Esparta, Costa Rica57.6-66.044.6-49.365.5-83.6(61.3)(47.3)(75.2)Ciudad Victoria, Tamaulipas50.6-56.944.5-48.767.2-84.3(53.7)(46.6)(73.9)Córdoba, Veracruz53.8-63.443.9-48.466.1-75.9(57.5)(45.6)(70.0)Isla del Carmen, Campeche47.3-56.644.7-48.961.5-72.6(50.9)(47.6)(65.7)Chichén-Itzá, Yucatán49.6-57.145.2-53.462.7-80.7(53.8)(49.5)(72.6)British Honduras49.0-59.647.5-50.767.9-76.8(54.9)(49.1)(72.2)Chinajá, Guatemala56.8-67.647.0-51.070.0-82.(63.2)(49.5)(73.6Atlantidad, Honduras52.5-65.149.8-53.656.1-76.5(57.6)(51.5)(67.0)Limón, Costa Rica57.7-71.350.4-52.363.9-73.0(62.4)(51.2)(68.5)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 thetympanum; 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 ofSmilisca baudiniis 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 21006from 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 ofSmilisca baudiniare 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 rangeSmilisca baudinioccurs 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 baudiniseeks 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 treesaround the water. Stuart (1935:38) recorded individuals calling and breeding throughout the day at La Libertad, Guatemala.Smilisca baudiniusually is absent from breeding congregations of hylids; frequentlyS. baudinibreeds alone in small temporary pools separated from large ponds where numerous other species are breeding. In Guerrero and Oaxaca, México,S. baudinibreeds in the same ponds withRhinophrynus dorsalis,Bufo marmoreus,Engystomops pustulosus, andDiaglena reticulata, and in the vicinity of Esparta, Costa Rica,S. baudinibreeds in ponds withBufo coccifer,Hyla staufferi, andPhrynohyas venulosa. In nearly all instances the breeding sites ofS. baudiniare shallow, temporary pools.The breeding call ofSmilisca baudiniconsists 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 patterndescribed 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 ofSmilisca baudinifrom 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 ofS. baudinifrom 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 ofSmiliscathat we have examined have a dextral anal tube.
University of Kansas PublicationsMuseum of Natural HistoryVolume 17, No. 7, pp. 281-375, pls. 1-12, 17 figs.July 14, 1966Neotropical Hylid Frogs, Genus SmiliscaBYWILLIAM E. DUELLMAN AND LINDA TRUEBUniversity of Kansas Lawrence1966
University of Kansas PublicationsMuseum 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 HistoryEditors: E. Raymond Hall, Chairman, Henry S. Fitch, Frank B. CrossVolume 17, No. 7, pp. 281-375, pls. 1-12, 17 figs.Published July 14, 1966University of KansasLawrence, KansasPRINTED BYROBERT R. (BOB) SANDERS, STATE PRINTERTOPEKA, KANSAS1966Look for the Union Label31-3430
University of Kansas Publications, Museum of Natural History
University of Kansas
PRINTED BYROBERT R. (BOB) SANDERS, STATE PRINTERTOPEKA, KANSAS1966Look for the Union Label31-3430
Look for the Union Label
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.Hylaand 10 other genera inhabit Central America; four of those 10 genera (Gastrotheca,Hemiphractus,Phrynohyas, andPhyllomedusa) are widely distributed in South America. The other six genera are either restricted to Central America or have their greatest differentiation there.PlectrohylaandPtychohylainhabit streams in the highlands of southern Mexico and northern Central America;DiaglenaandTriprionare casque-headed inhabitants of arid regions in México and northern Central America.Anothecais a tree-hole breeder in cloud forests in Middle America. The genusSmiliscais 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 generaDiaglena,Triprion,Tetraprion,Osteocephalus,Trachycephalus,Aparasphenodon,Corythomantis,Hemiphractus,Pternohyla, andAnothecahave been recognized as distinct from one another and from the genusHylaon the basis of various modifications of dermal bones of the cranium.Phyllomedusais recognized on the basis of a vertical pupil and opposable thumb;Plectrohylais characterized by the presence of a bony prepollex and the absence of a quadratojugal.Gastrothecais 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 distinguishPhrynohyasfromHyla. He (1963a) cited the ventrolateral glands in breeding males as diagnostic ofPtychohyla. Some species groups within the vaguely defined genusHylahave equally distinctive characters. TheHyla septentrionalisgroup is characterized by a casque-head, not much different from that in the genusOsteocephalus(Trueb, MS). Males in theHyla maximagroup have a protruding bony prepollex like that characteristically found inPlectrohyla.
Ontogenetic development, osteology, breeding call, behavior, and ecology are important in the recognition of species. By utilizingthe combination of many morphological and biological factors, the genusSmiliscacan be defined reasonably well as a natural, phyletic assemblage of species. Because the wealth of data pertaining to the morphology and biology ofSmiliscais lacking for most other tree frogs in Middle America it is not possible at present to compareSmiliscawith related groups in more than a general way.
Smiliscais 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 genusSmilisca, 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 genusSmiliscato 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.
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 andcleanedand stained specimens of all species, plus serial sections of the skull ofSmilisca 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 SmiliscaCope, 1865
SmiliscaCope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct., 1865 [Type speciesSmilisca dauliniaCope, 1865 =Hyla baudiniDuméril 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 mandibulaeconsisting of two parts, one arising from dorsal fascia and other from posterior arm of squamosal, (6) dividedM. 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 genusSmiliscacontains six recognizable species. An alphabetical list of the specific and subspecific namesthat we consider to be applicable to species ofSmiliscarecognized 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 baudiniDuméril 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 vanvlietiiBaird, 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 vociferansBaird, 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 muricolorCope, 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 dauliniaCope, 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 forbaudini], 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 pansosanaBrocchi, 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 beltraniTaylor, 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 manisorumTaylor, 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 distinguishingS. baudinifrom 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 thetympanum; 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 ofSmilisca baudiniis 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 21006from 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 ofSmilisca baudiniare 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 rangeSmilisca baudinioccurs 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 baudiniseeks 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 treesaround the water. Stuart (1935:38) recorded individuals calling and breeding throughout the day at La Libertad, Guatemala.Smilisca baudiniusually is absent from breeding congregations of hylids; frequentlyS. baudinibreeds alone in small temporary pools separated from large ponds where numerous other species are breeding. In Guerrero and Oaxaca, México,S. baudinibreeds in the same ponds withRhinophrynus dorsalis,Bufo marmoreus,Engystomops pustulosus, andDiaglena reticulata, and in the vicinity of Esparta, Costa Rica,S. baudinibreeds in ponds withBufo coccifer,Hyla staufferi, andPhrynohyas venulosa. In nearly all instances the breeding sites ofS. baudiniare shallow, temporary pools.
The breeding call ofSmilisca baudiniconsists 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 patterndescribed 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 ofSmilisca baudinifrom 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 ofS. baudinifrom 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 ofSmiliscathat we have examined have a dextral anal tube.