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diff --git a/37823.txt b/37823.txt new file mode 100644 index 0000000..a57ac86 --- /dev/null +++ b/37823.txt @@ -0,0 +1,6224 @@ +The Project Gutenberg EBook of Neotropical Hylid Frogs, Genus Smilisca, by +William E. Duellman and Linda Trueb + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Neotropical Hylid Frogs, Genus Smilisca + +Author: William E. Duellman + Linda Trueb + +Release Date: October 22, 2011 [EBook #37823] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK NEOTROPICAL HYLID FROGS *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + + Volume 17, No. 7, pp. 281-375, pls. 1-12, 17 figs. + + July 14, 1966 + + + Neotropical Hylid Frogs, Genus Smilisca + + BY + + WILLIAM E. DUELLMAN AND LINDA TRUEB + + + UNIVERSITY OF KANSAS + LAWRENCE + 1966 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Henry S. Fitch, + Frank B. Cross + + + Volume 17, No. 7, pp. 281-375, pls. 1-12, 17 figs. + Published July 14, 1966 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY + ROBERT R. (BOB) SANDERS, STATE PRINTER + TOPEKA, KANSAS + 1966 + + [Illustration: Look for the Union label.] + + 31-3430 + + + + + Neotropical Hylid Frogs, Genus Smilisca + + + BY + + + WILLIAM E. DUELLMAN AND LINDA TRUEB + + + + + CONTENTS + + + PAGE + + INTRODUCTION 285 + Acknowledgments 286 + Materials and Methods 287 + + GENUS SMILISCA COPE, 1865 287 + Key to Adults 288 + Key to Tadpoles 289 + + ACCOUNTS OF SPECIES 289 + _Smilisca baudini_ (Dumeril and Bibron) 289 + _Smilisca cyanosticta_ (Smith) 303 + _Smilisca phaeota_ (Cope) 308 + _Smilisca puma_ (Cope) 314 + _Smilisca sila_ New species 318 + _Smilisca sordida_ (Peters) 323 + + ANALYSIS OF MORPHOLOGICAL CHARACTERS 330 + Osteology 330 + _Descriptive Osteology of Smilisca baudini_ 331 + _Developmental Cranial Osteology of Smilisca baudini_ 333 + _Comparative Osteology_ 336 + Musculature 341 + Skin 342 + _Structure_ 342 + _Comparative Biochemistry of Proteins_ 343 + External Morphological Characters 343 + _Size and Proportions_ 343 + _Shape of Snout_ 344 + _Hands and Feet_ 344 + _Ontogenetic Changes_ 344 + Coloration 344 + _Metachrosis_ 345 + Chromosomes 345 + + NATURAL HISTORY 345 + Breeding 345 + _Time of Breeding_ 345 + _Breeding Sites_ 346 + _Breeding Behavior_ 346 + _Breeding Call_ 351 + Eggs 356 + Tadpoles 357 + _General Structure_ 357 + _Comparison of Species_ 357 + _Growth and Development_ 361 + _Behavior_ 365 + + PHYLOGENETIC RELATIONSHIPS 366 + Interspecific Relationships 366 + Evolutionary History 369 + + SUMMARY AND CONCLUSIONS 371 + + LITERATURE CITED 372 + + + + +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 Mexico 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, Guenther 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 Jose, 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 Mexico, 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 Panama under the +auspices of a grant from the National Institutes of Health (NIH +GM-12020) in cooperation with the Gorgas Memorial Laboratory in Panama. + + +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 Mexico 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: + + AMNH--American Museum of Natural History + BMNH--British Museum (Natural History) + BYU--Brigham Young University + CNHM--Chicago Natural History Museum + KU--University of Kansas Museum of Natural History + MCZ--Museum of Comparative Zoology + MNHN--Museu National d'Histoire Naturelle, Paris + UF--University of Florida Collections + UIMNH--University of Illinois Museum of Natural History + UMMZ--University of Michigan Museum of Zoology + USC--University of Southern California + USNM--United States National Museum + TNHC--Texas Natural History Collection, University of Texas + ZMB--Zoologisches Museum Berlin + + +=Genus Smilisca= Cope, 1865 + + _Smilisca_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct., + 1865 [Type species _Smilisca daulinia_ Cope, 1865 = _Hyla baudini_ + Dumeril 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. + + + Names proposed Valid names + + _Hyla baudini_ Dumeril and Bibron, 1841 = _S. baudini_ + _Hyla baudini dolomedes_ Barbour, 1923 = _S. phaeota_ + _Hyla beltrani_ Taylor, 1942 = _S. baudini_ + _Hyla gabbi_ Cope, 1876 = _S. sordida_ + _Hyla labialis_ Peters, 1863 = _S. phaeota_ + _Hyla manisorum_ Taylor, 1954 = _S. baudini_ + _Hyla muricolor_ Cope, 1862 = _S. baudini_ + _Hyla nigripes_ Cope, 1876 = _S. sordida_ + _Hyla pansosana_ Brocchi, 1877 = _S. baudini_ + _Hyla phaeota_ Cope, 1862 = _S. phaeota_ + _Hyla phaeota cyanosticta_ Smith, 1953 = _S. cyanosticta_ + _Hyla puma_ Cope, 1885 = _S. puma_ + _Hyla salvini_ Boulenger, 1882 = _S. sordida_ + _Hyla sordida_ Peters, 1863 = _S. sordida_ + _Hyla vanvlietii_ Baird, 1854 = _S. baudini_ + _Hyla vociferans_ Baird, 1859 = _S. baudini_ + _Hyla wellmanorum_ Taylor, 1952 = _S. puma_ + + +_Distribution of genus._--Most of lowlands of Mexico and Central +America, in some places to elevations of nearly 2000 meters, southward +from southern Sonora and Rio Grande Embayment of Texas, including such +continental islands as Isla Cozumel, Mexico, and Isla Popa and Isla +Cebaco, Panama, to northern South America, where known from Caribbean +coastal regions and valleys of Rio Cauca and Rio Magdalena in Colombia, +and Pacific slopes of Colombia and northern Ecuador. + + +Key to Adults + + 1. Larger frogs ([M] to 76 mm., [F] to 90 mm.) having broad + flat heads and a dark brown or black postorbital mark + encompassing tympanum 2 + + Smaller frogs ([M] to 45 mm., [F] to 64 mm.) having narrower + heads and lacking a dark brown or black postorbital mark + encompassing tympanum 4 + + 2. Lips barred; flanks cream-colored with bold brown or black + mottling in groin; posterior surfaces of thighs brown with + cream-colored flecks _S. baudini_, p. 289 + + Lips not barred; narrow white labial stripe present; flanks + not cream-colored with bold brown or black mottling in groin; + posterior surfaces of thighs variable 3 + + 3. Flanks and anterior and posterior surfaces of thighs dark + brown with large pale blue spots on flanks and small blue + spots on thighs _S. cyanosticta_, p. 303 + + Flanks cream-colored with fine black venation; posterior + surfaces of thighs pale brown with or without darker flecks + or small cream-colored spots _S. phaeota_, p. 308 + + 4. Fingers having only vestige of web; diameter of tympanum + two-thirds that of eye; dorsum pale yellowish tan with pair + of broad dark brown stripes _S. puma_, p. 314 + + Fingers about one-half webbed; diameter of tympanum about + one-half that of eye; dorsum variously marked with spots + or blotches 5 + + 5. Snout short, truncate; vocal sacs in breeding males dark + gray or brown; blue spots on flanks and posterior surfaces + of thighs _S. sila_, p. 318 + + Snout long, sloping, rounded; vocal sacs in breeding males + white; cream-colored or pale blue flecks on flanks and + posterior surfaces of thighs _S. sordida_, p. 323 + + +Key to Tadpoles + + 1. Pond tadpoles; tail about half again as long as body; + mouth anteroventral 2 + + Stream tadpoles; tail about twice as long as body; + mouth ventral 5 + + 2. Labial papillae in two rows 3 + + Labial papillae in one row 4 + + 3. 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 reticulations _S. puma_, p. 314 + + First 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 + blotches _S. baudini_, p. 289 + + 4. Dorsal fin extending onto body _S. phaeota_, p. 308 + + Dorsal fin not extending onto body _S. cyanosticta_, p. 303 + + 5. 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 tail + _S. sordida_, p. 323 + + Median 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 tail _S. sila_, p. 318 + + + + +ACCOUNTS OF SPECIES + + +=Smilisca baudini= (Dumeril and Bibron) + + _Hyla baudini_ Dumeril and Bibron, Erpetologie general, 8:564, 1841 + [Holotype.--MNHN 4798 from "Mexico;" Baudin collector]. Guenther, + Catalogue Batrachia Salientia in British Museum, p. 105, 1858. + Brocchi, Mission scientifique au Mexique ..., pt. 3, sec. 2, Etudes + 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. Guenther, 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 Mexico, 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_]. Diaz de Leon, Indice de los batracios + que se encuentran en la Republica Mexicana, p. 20, June 1904. + + _Hyla muricolor_ Cope, Proc. Acad. Nat. Sci. Philadelphia, + 14(9):359, 1862 [Holotype.--USNM 25097 from Mirador, Veracruz, + Mexico; 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 Panzos, Alta Verapaz, Guatemala; M. + Bocourt collector]; Mission scientifique au Mexique ..., pt. 3, + sec. 2, Etudes 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, Mexico; A. Magana 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 Batan, Limon 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 Mexico, including Tamaulipas, Veracruz, the Yucatan +Peninsula, and British Honduras. + + + Table 1.--Geographic Variation in Size and Proportions in Males of + Smilisca baudini. (Means in Parentheses Below Observed Ranges; + Data Based on 10 Specimens From Each Locality.) + + ============================+============+==============+============ + Locality | Snout-vent | Tibia length/| Tympanum/ + | length | snout-vent | eye + ----------------------------+------------+--------------+------------ + Southern Sinaloa | 62.3-75.9 | 43.2-46.7 | 84.2-94.4 + | (68.6) | (44.9) | (87.8) + | | | + Ocotito, Guerrero | 55.6-64.0 | 46.1-51.2 | 66.7-82.8 + | (58.7) | (47.8) | (74.6) + | | | + Pochutla, Oaxaca | 56.1-65.1 | 44.7-49.4 | 73.0-84.2 + | (60.2) | (47.5) | (77.4) + | | | + San Salvador, El Salvador | 57.0-68.0 | 42.1-46.1 | 74.6-83.3 + | (62.1) | (44.9) | (77.6) + | | | + Managua, Nicaragua | 52.9-63.6 | 45.6-49.4 | 73.7-89.7 + | (57.3) | (47.5) | (79.4) + | | | + Esparta, Costa Rica | 57.6-66.0 | 44.6-49.3 | 65.5-83.6 + | (61.3) | (47.3) | (75.2) + | | | + Ciudad Victoria, Tamaulipas | 50.6-56.9 | 44.5-48.7 | 67.2-84.3 + | (53.7) | (46.6) | (73.9) + | | | + Cordoba, Veracruz | 53.8-63.4 | 43.9-48.4 | 66.1-75.9 + | (57.5) | (45.6) | (70.0) + | | | + Isla del Carmen, Campeche | 47.3-56.6 | 44.7-48.9 | 61.5-72.6 + | (50.9) | (47.6) | (65.7) + | | | + Chichen-Itza, Yucatan | 49.6-57.1 | 45.2-53.4 | 62.7-80.7 + | (53.8) | (49.5) | (72.6) + | | | + British Honduras | 49.0-59.6 | 47.5-50.7 | 67.9-76.8 + | (54.9) | (49.1) | (72.2) + | | | + Chinaja, Guatemala | 56.8-67.6 | 47.0-51.0 | 70.0-82.8 + | (63.2) | (49.5) | (73.6) + | | | + Atlantidad, Honduras | 52.5-65.1 | 49.8-53.6 | 56.1-76.5 + | (57.6) | (51.5) | (67.0) + | | | + Limon, Costa Rica | 57.7-71.3 | 50.4-52.3 | 63.9-73.0 + | (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 Mexico the average ratio varies from +45.6 in Veracruz to 47.6 on Isla del Carmen, Campeche. Specimens from +the Yucatan 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 Limon, 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 Chinaja, +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 Mexico, +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 Matatan, 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 +Coban 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 Limon Province, Costa Rica (KU 34927 from +Batan 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 Peten +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 Mexico 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 +Peten, 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 Peten, +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 Chinaja, 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, Mexico, _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 Chinaja, +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, Yucatan (KU 71720), and +4 kilometers west-southwest Puerto Juarez, Quintana Roo, Mexico (KU +71721), whereas others, notably from 17 kilometers northeast of +Juchatengo, Oaxaca, Mexico (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 Piste, Yucatan, Mexico. These +specimens are heavily pigmented like specimens that we have examined +from the Yucatan 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 Peten, 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, Mexico, 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. Dumeril and Bibron (1841:564) proposed the name +_Hyla baudini_ for a specimen (MNHN 4798) from Mexico. Smith and Taylor +(1950:347) restricted the type locality to Cordoba, Veracruz, Mexico, 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, Mexico, 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. + + + [Illustration: PLATE 1 + + A + + B + + Dorsal views of skulls of young _Smilisca baudini_: (A) recently + metamorphosed young (KU 60026), snout-vent length 12.6 mm. x23; + (B) young (KU 85438), snout-vent length 32.1 mm. x9.] + + [Illustration: PLATE 2 + + A + + B + + Skull of adult female _Smilisca baudini_ (KU 68184): (A) Dorsal; + (B) Ventral. x4.5.] + + [Illustration: PLATE 3 + + A + + B + + C + + Skull of adult female _Smilisca baudini_ (KU 68184): (A) Lateral; + (B) Dorsal view of left mandible; (C) Posterior. x4.5.] + + [Illustration: PLATE 4 + + A B + + C D + + E F + + 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). x3.] + + [Illustration: PLATE 5 + + A B + + C D + + E F + + 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). x3.] + + [Illustration: PLATE 6 + + A + + B + + C + + Living _Smilisca_: (A) _S. baudini_ (UMMZ 115179) from 1.7 km. W + Xicotencatl, Tamaulipas, Mexico; (B) _S. cyanosticta_ (UMMZ 118163) + from Volcan San Martin, Veracruz, Mexico; (C) _S. phaeota_ + (KU 64282) from Barranca del Rio Sarapiqui, Heredia Prov., Costa Rica. + All approx. nat. size.] + + [Illustration: PLATE 7 + + A + + B + + C + + 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 Volcan, Chiriqui, Panama; + (C) _S. sordida_ (KU 64257) from 20 km. WSW San Isidro el General, + San Jose Prov., Costa Rica. All approx. nat. size.] + + [Illustration: 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.] + + [Illustration: 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_, Rio La Vieja, 30 km. + E of Palmar Norte, Puntarenas Prov., Costa Rica.] + + [Illustration: 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).] + + [Illustration: 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).] + + [Illustration: 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). x3.2.] + +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 Batan, +Limon, 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 Rio +Esnape, Darien, Panama. + + [Illustration: 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 +Mexico from southern Sonora and the Rio 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 Solistahuacan, Chiapas, elevation 1675 meters, +and 10 kilometers northwest of Comitan, 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; Champoton, UMMZ 73172 (2), 73176, 73180; 16 km. E +Champoton, UMMZ 73181; 5 km. S Champoton, KU 71369-75; 9 km. S +Champoton, KU 71367-8; 10.5 km. S Champoton, KU 71365-6, 71722 +(tadpoles), 71723 (yg.); 24 km. S Champoton, UMMZ 73177 (2); Chuina, KU +75101-3; Ciudad del Carmen, UIMNH 30703-8; Dzibalchen, KU 75413-31; +Encarnacion, CNHM 102282, 102289, 102294-5, 102300, 102306-8, 102312, +102314, 102316-7, 102319, 102322, UIMNH 30727-40, 30836-7; 1 km. W +Escarcega, KU 71391-6; 6 km. W Escarcega, KU 71397-403; 7.5 km. W +Escarcega, KU 71376-89; 14 km. W Escarcega, KU 71390; 13 km. W, 1 km. N +Escarcega, KU 71404; 3 km. N Hopelchen, KU 75410-11; 2 km. NE Hopelchen, +KU 75412; Matamoras, CNHM 36573; Pital, UIMNH 30741; 1 km. SW Puerto +Real, Isla del Carmen, KU 71345-64; San Jose Carpizo, UMMZ 99879; Tres +Brazos, CNHM 102284, UIMNH 30723-5; Tuxpena 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); +Comitan, UMMZ 94438; 10 km. NW Comitan, 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 Juarez, 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 +Solistahuacan, 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 Geronimo, 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; Tonola, 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 Rio, +CNHM 102207, 102229-30, UIMNH 30819-21, UMMZ 110875 (3); Periquillo, +UMMZ 80025 (3), 80026 (14); 1.6 km. SW Pueblo Juarez, 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 Mazatlan, UIMNH 30777-9; 9 km. S Mazatlan, 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 Marquez, AMNH 59200-1 (13); +5.6 km. S San Andreas de la Cruz, KU 87173-4; San Vincente, KU 87172; +Zaculapan, UMMZ 119183. + +=Hidalgo=: Below Tianguistengo, CNHM 102318. + +=Jalisco=: Atenqueque, KU 91435-6; 5 km. NE Autlan, 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. + +=Michoacan=: Aguililla, UMMZ 119179 (5); Apatzingan, CNHM 38766-90, KU +69101 (skeleton); 7 km. E Apatzingan, UMMZ 112843; 11 km. E Apatzingan, +UMMZ 112841 (3); 27 km. S Apatzingan, 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 Jose de la Montana, 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; Jesus Maria, 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 Leon=: 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); Juchitan, USNM 70400; Lagartero, UIMNH 9514; +Matias Romero, AMNH 52143-5; 25 km. N Matias Romero, KU 33822-8; 7 km. S +Matias Romero, UIMNH 42703; Mirador, AMNH 6277, 13832-9, 13842-55; Mira +Leon, 1.6 km. N Huatulco, UIMNH 9503-4; Mixtequillo, AMNH 13924; +Pochutla, KU 57167-81, UIMNH 9505-13; Quiengola, AMNH 51817, 52146; Rio +del Corte, UIMNH 48677; Rio Mono Blanco, UIMNH 36831; Rio 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 Juarez, UF 11205. + +=Quintana Roo=: Coba, 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 Juarez, KU 71407-11, +71721 (tadpoles); 12 km. W Puerto Juarez, 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 Potosi=: 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); Rio 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 Cosala, KU 73910; Costa Rica, +16 km. S. Culiacan, UIMNH 34887-9; 51 km. SSE Culiacan, 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; +Matatan, KU 73913; 7.3 km. SW Matatan, KU 78464, 78466-70; Mazatlan, +AMNH 12562, UMMZ 115197 (3); 57 km. N Mazatlan, 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; Teacapan, Isla +Palmito del Verde, KU 73915; 9.6 km. NNW Teacapan, KU 91410; Villa +Union, KU 78471; 9 km. NE Villa Union, KU 75434-7; 1 km. W Villa Union, +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=: Acuna, UMMZ 99864; 5 km. S Acuna, UMMZ 101180; 13 km. N +Antiguo Morelos, UIMNH 40532-5; 3 km. S Antiguo Morelos, UF 11204; 3 km. +NE Chamal, UMMZ 102867; 1.6 km. E Chamal, UMMZ 110734; Ciudad Mante, +UMMZ 80957, 80958 (3), 106400 (3); 16 km. N Ciudad Victoria, CNHM +102408; 34 km. NE Ciudad Victoria, KU 60395-411; 8.8 km. S Ciudad +Victoria, UIMNH 19261-3; 11 km. W Ciudad Victoria, UIMNH 30924; 16 km. W +Ciudad Victoria, UIMNH 30825; 3 km. W El Carizo, UMMZ 111279; Gomez +Farias, UMMZ 110837-8; 8 km. NE Gomez Farias, UMMZ 102265, 102916 (4), +102917, 104110 (5), 105493, 110836 (2), 111274-7; 8 km. NW Gomez Farias, +UMMZ 101178 (7), 101179 (3), 101362-3, 101364 (2), 108799 (2), 110129, +111278, 111280; 8 km. W Gomez Farias, UMMZ 102859 (2); 16 km. W +Gonzales, KU 37795-6; Jimenez, KU 60412; La Clementina, 6 km. W Forlan, +USNM 106244; Limon, UIMNH 30831; Llera, USNM 140137-40; 3 km. E Llera, +UIMNH 16858; 21 km. S Llera, UIMNH 30828-9; 23 km. S Llera, UIMNH 30830; +11 km. SW Ocampo, UMMZ 118956; 22 km. W, 5 km. S Piedra, KU 37568-71; +Rio Sabinas, UMMZ 97976; 5 km. W San Gerardo, UMMZ 110733 (2); Santa +Barbara, UMMZ 111272-3; Villagran, CNHM 102280, 102287, 102299, 102309, +UIMNH 30826-7; 1.7 km. W Xicotencatl, UMMZ 115179. + +=Veracruz=: 1.6 km. NW Acayucan, UMMZ 115189; 28.5 km. SE Alvarado, UMMZ +119933; 2.4 km. SSW Amatitlan, UMMZ 115195; Barranca Metlac, UIMNH +38365; Boca del Rio, UIMNH 26619-30, UMMZ 74954 (9); 16 km. S Boca del +Rio, UIMNH 26631; between Boca del Rio and Anton Lizardo, UIMNH 42701; +Canada, CNHM 102397; Catemaco, UMMZ 118702 (4); Ciudad Aleman, UMMZ +119608 (3); Cordoba, CNHM 38665-7, USNM 30410-3; 5.2 km. ESE Cordoba, KU +71423-35, 89924 (skeleton); 7 km. ESE Cordoba, UMMZ 115176 (4); +Cosamaloapan, UMMZ 115193 (2); Coyame, UIMNH 36853-6, 38366, UMMZ 111461 +(3), 111462-3; 1 km. SE Coyame, UMMZ 121202 (3); Cuatotolapam, UMMZ +41625-39; Cuautlapan, CNHM 38664, 70591-600, 102218, 102398, KU 26300, +26302, 26309, 26312-3, 26315-6, 26321, 26336, 26339, 26347 (skeleton), +26354, 55614-21 (skeletons), UIMNH 11236-67, 11269-71, 11273, 26611-8, +30792-5, UMMZ 85466 (6), 115173 (25), 115175 (7), USNM 114433-57; Dos +Rios, CNHM 39623; 5 km. ENE El Jobo, KU 23843, 23845, 23847; 6.2 km. E +Encero, UIMNH 30835; Escamilo, CNHM 102298, UIMNH 30788; 1 km. N Fortin, +UF 11201; 4 km. SW Huatusco, UMMZ 115177; 1 km. SW Huatusco, UMMZ +123119; 10 km. SE Hueyapan, UMMZ 115190; 20 km. S Jesus Carranza, KU +23844, 23846, 27399; 38 km. SE Jesus Carranza, KU 23417; Laguna +Catemaco, UMMZ 119932 (62); 1.6 km. N La Laja, UIMNH 3651; La Oaxaquena, +AMNH 43930-1; 17 km. E Martinez de la Torre, UIMNH 36630-2; 6.2 km. W +Martinez de la Torre, UIMNH 3652-4; Minatitlan, AMNH 52141-2; Mirador, +USNM 25097-8, 115178; 6 km. S Monte Blanco, UF 11200 (4); 21 km. E +Nanchital, UMMZ 123004; 2 km. S Naranja, UMMZ 115188 (3); 1.6 km. NE +Novillero, UMMZ 115194 (2); 3 km. NE Novillero, UMMZ 115196; 5.2 km. NE +Novillero, UMMZ 115192 (4); 6 km. NE Novillero, UMMZ 115191; 5 km. N +Nuevo Colonia, UMMZ 105066; Orizaba, USNM 16563-6; 4 km. NE Orizaba, +UMMZ 120251 (2); Panuco, UMMZ 118922; Paraje Nuevo, UMMZ 85465 (2), +85467 (35), 85468 (36); Paso del Macho, UIMNH 49281; Paso de Talaya, +Jicaltepec, USNM 32365, 84420; Perez, CNHM 1686 (5); 20 km. N Piedras +Negras, Rio Blanco, KU 23708; Plan del Rio, KU 26310, 26333-5, 26345, +26354, UMMZ 102069, 102070 (5); Potrero, UIMNH 49282-5, UMMZ 88799, +88805, 88806 (2), USNM 32391-5; Potrero Viejo, CNHM 102296, KU 26301, +26304-5, 26307-8, 26311, 26317-20, 26323-25, 26326-8 (skeletons), +26329-31, 26332 (skeleton), 26337-8, 26340-4, 26346, 26348, 26351, +26353, 27400-12, UIMNH 30800, UMMZ 88800 (2), 88802 (15), 88803 (9), +88804, USNM 114458-67; 5 km. S Potrero Viejo, KU 26303, 26314, 26322; +Puente Nacional, UIMNH 21783-8; 3 km. N Rinconada, UMMZ 122099 (5); Rio +de las Playas, USNM 118635-6; Rio Seco, UMMZ 88801 (9); Rodriguez Clara, +CNHM 102225; San Andres Tuxtla, CNHM 102213, 102222, 102227, 102247, +UIMNH 30789-91; 10 km. NW San Andres Tuxtla, UMMZ 119935; 13.4 km. NW +San Andres Tuxtla, UMMZ 119939 (2); 19.8 km. NW San Andres Tuxtla, UMMZ +119938; 27.2 km. NW San Andres Tuxtla, UMMZ 119936 (6); 48 km. NW San +Andres Tuxtla, UMMZ 119937; 4 km. W San Andres Tuxtla, UMMZ 115187; 37.4 +km. S San Andres Tuxtla, UMMZ 119934 (12); 15 km. ESE San Juan de la +Punta, KU 23707; San Lorenzo, USNM 123508-12; 3 km. SW San Marcias KU +23841; 1.5 km. S Santa Rosa, UIMNH 42702; 2 km. S Santiago Tuxtla, UMMZ +121201 (4); Sauzel, UMMZ 121239; 14 km. E Suchil, UIMNH 46880; 15 km. S +Tampico (Tamaulipas), UMMZ 103322 (4); 4 km. N Tapalapan, UMMZ 115186 +(2); Tecolutla, UIMNH 42677-700; 16 km. NW Tehuatlan, UIMNH 3660-3; +5 km. S Tehuatlan, KU 23842; Teocelo, KU 26306; Tierra Colorado, CNHM +102393, 102395-6, UIMNH 30789-91; Veracruz, AMNH 6301-4, 59398-402, +UIMNH 30801, UMMZ 115174, 122060 (2); 24 km. W Veracruz, CNHM 104570-2. + +=Yucatan=: No specific locality, CNHM 548, 49067, USNM 32298; +Chichen-Itza, CNHM 20636, 26938-49, 36559-62, UIMNH 30742-6, UMMZ 73173 +(6), 73174 (14), 73175 (14), 73178-9, 76171, 83107 (2), 83108, 83109 +(2), 83915 (30), USNM 72744; 9 km. E Chichen-Itza, KU 71438-9; 12 km. E +Chichen-Itza, KU 71440; Merida, CNHM 40659-66, UIMNH 30747-8, UMMZ +73182; 6 km. S Merida, KU 75194; 8.8 km. SE Ticul, UMMZ 114296; +Valladolid, CNHM 26934-6; Xcalah-op, CNHM 53906-14; 3.5 km. E Yokdzonot, +KU 71441-3, 71720 (tadpoles). + +BRITISH HONDURAS: Belize, CNHM 4153, 4384-5, 4387, UMMZ 75310, USNM +26065; Bokowina, CNHM 49064-5; Cocquercote, UMMZ 75331 (2); Cohune +Ridge, UMMZ 80738 (15); Double Falls, CNHM 49066; El Cayo, UMMZ 75311; +6 km. S El Cayo, MCZ 37856; Gallon Jug, MCZ 37848-55; Manatee, CNHM +4264-7; Mountain Pine Ridge, MCZ 37857-8; San Augustin, UMMZ 80739; San +Pedro, Columbia, MCZ 37860-2; Valentin, UMMZ 80735 (4), 80736 (2), 80737 +(2); 5 km. S Waha Loaf Creek, MCZ 37859. + +GUATEMALA: =Alta Verapaz=: 5.1 km. NE Campur, KU 68464 (tadpoles), 67465 +(young); 28.3 km. NE Campur, KU 64203-22, 68183-4 (skeletons); Chama, +MCZ 15792-3, UMMZ 90895 (7), 90896 (5), 90897 (29), 90898 (12), 90899; +Chinaja, KU 55939-41, 57193-8, 60018-20 (tadpoles), 60021 (eggs), 60022 +(tadpoles); Coban, CNHM 21006; Cubilquitz, UMMZ 90902 (10); Finca +Canihor, UMMZ 90908; Finca Chicoyou, KU 57246-8, 60026 (young), 64202, +68466-7 (tadpoles); Finca Los Alpes, KU 64197-201, 68463 (tadpoles); +Finca Los Pinales, UMMZ 90903 (2); Finca Tinajas, BYU 16031; Finca +Volcan, UMMZ 90905 (4), 90906-7; Panzos, MNHN 6313, UMMZ 90904; Samac, +UMMZ 90900; Samanzana, UMMZ 90901 (6). + +=Baja Verapaz=: Chejel, UMMZ 90909 (7), 90910 (3); San Geronimo, UMMZ +84076 (16). + +=Chiquimula=: 1.6 km. SE Chiquimula, UMMZ 98112; Esquipulas, UMMZ +106793 (28). + +=El Peten=: 20 km. NNW Chinaja (Alta Verapaz), KU 57199-240; Flores, +UMMZ 117985; La Libertad, KU 60024 (young), UMMZ 75313-20, 75323 (2), +75324 (7), 75325 (13), 75326 (2), 75327 (11), 75328 (12), 75329 (2); +3 km. SE La Libertad, KU 57243-4; 13 km. S La Libertad, MCZ 21458 (2); +Pacomon, USNM 71334; Piedras Negras, USNM 114469-71; Poptun, UMMZ +120475; Poza de la Jicotea, USNM 114672; Ramate-Yaxha trail, UMMZ 75321; +Rio de la Pasion between Sayaxche and Subin, KU 57151; Rio San Roman, +16 km. NNW Chinaja (Alta Verapaz), KU 55942-6; Sacluc, USNM 25131; +Sayaxche, KU 57144-5; Tikal, UMMZ 117983 (7), 117984 (5), 117993 (5), +120474 (5); Toocog, KU 57241-2, 60023 (young), 60025 (young); Uaxactun, +UMMZ 70401-3; Yaxha, UMMZ 75322; 19 km. E Yaxha, UMMZ 75330 (4). + +=El Quiche=: Finca Tesoro, UMMZ 89166 (3), 90549 (tadpoles). + +=Escuintla=: Rio Guacalate, Masagua, USNM 125239; Tiquisate, +UMMZ 98262 (7). + +=Guatemala=: 16 km. NE Guatemala, KU 43545-53. + +=Huehuetenango=: Finca San Rafael, 16 km. SE Barillas, CNHM 40912-6; +45 km. WNW Huehuetenango, KU 64223-4; Jacaltenango, UMMZ 120080 (6), +120081 (14), 120082 (13). + +=Izabal=: 2 km. SW Puerto Matias de Galvez, KU 60027-8 (tadpoles); +Quirigua, CNHM 20587, UMMZ 70060. + +=Jalapa=: Jalapa, UMMZ 98109, 106792 (11). + +=Jutiapa=: Finca La Trinidad, UMMZ 107728 (10); Jutiapa, UMMZ 106789; +1.6 km. SE Mongoy, KU 43069; Santa Catarina Mita, UMMZ 106790. + +=Progreso=: Finca Los Leones, UMMZ 106791. + +=Quetzaltenango=: Coatepeque, AMNH 62204. + +=Retalhueleu=: Casa Blanca, UMMZ 107725 (18); Champerico, UMMZ 107726 +(3). + +=San Marcos=: Talisman Bridge, USNM 128056-7. + +=Santa Rosa=: Finca La Guardiana, UMMZ 107727 (6); Finca La Gloria, UMMZ +107724 (6); 1.6 km. WSW El Molino, KU 43065-8. + +EL SALVADOR: =La Libertad=: 16 km. NW Santa Tecla, KU 43542-4. +=Morazan=: Divisadero, USNM 73284. =San Salvador=: San Salvador, CNHM +65087-99, KU 61955-88, 62138-9 (skeletons), 62154 (eggs), 62155-60 +(tadpoles), 68462 (tadpoles), UMMZ 117586 (3), 118380 (3), USNM 140278. + +HONDURAS: State unknown: Guaimas, UMMZ 58391. =Atlantidad=: Isla de +Roatan, CNHM 34551-4; La Ceiba, USNM 64985, 117589-91; Lancetilla, MCZ +16207-11; Tela, MCZ 15774-5, 28080, UMMZ 58418, USNM 82173-4. +=Choluteca=: 1.5 km. NW Choluteca, KU 64228-32; 10 km. NW Choluteca, KU +64233; 10 km. E Choluteca, KU 64226-7; 12 km. E Choluteca, KU 64225; 5 +km. S Choluteca, USC 2700 (2). =Colon=: Bambu, UF 320; Belfate, AMNH +45692-5; Patuca, USNM 20261. =Comayagua=: La Mision, 3.5 leagues N +Siguatepeque, MCZ 26424-5. =Copan=: Copan, UMMZ 83026 (2). =Cortes=: +Cofradia, AMNH 45345-6; Hacienda Santa Ana, CNHM 4724-31; Lago de Yojoa, +MCZ 26410-1; Rio Lindo, AMNH 54972. =El Paraiso=: El Volcan, MCZ 26436. +=Francisco Morazan=: Tegucigalapa, BYU 18301-4, 18837-41, MCZ 26395-7, +USNM 60499. =Gracias A Dios=: Rio Segovia, MCZ 24543. =Santa Barbara=: +Santa Barbara, USNM 128062-5. + +NICARAGUA: =Chinandega=: 4 km. N, 2 km. W Chichigalpa, KU 85385; +Chinandega, MCZ 2632; Rio Tama, USNM 40022; San Antonio, KU 84944-9 +(skeletons), 85386-403. =Chontales=: 1 km. NE Acoyapa, KU 64234. +=Esteli=: Finca Daraili, 5 km. N, 15 km. E Condega, KU 85404-8; Finca +Venecia, 7 km. N, 16 km. E Condega, KU 85409. =Leon=: 1.6 km. ENE +Poneloya, KU 43084-5. =Managua=: Managua, USNM 79989-90; 8 km. NW +Managua, KU 43094-110; 20 km. NE Managua, KU 85412; 21 km. NE Managua, +KU 85413-4; 5 km. SW Managua, KU 43086-93; 2 km. N Sabana Grande, KU +85411; 3 km. N Sabana Grande, KU 43070-8; 20 km. S, 0.5 km. W Tipitapa, +KU 85410. =Matagalpa=: Guasqualie, UMMZ 116493; Matagalpa, UMMZ 116492; +19 km. N Matagalpa, UMMZ 116494. =Rio San Juan=: Greytown, USNM 19585-6, +19767-8. =Rivas=: Javillo, UMMZ 123001; Moyogalpa, Isla Ometepe, KU +85428-37, 87706 (tadpoles); Penas Blancas, KU 85417; Rio Javillo, 3 km. +N, 4 km. W Sapoa, KU 85418-20, 85438 (skeleton); 13.1 km. SE Rivas, KU +85415; 14.8 km. SE Rivas, KU 85421-3; 11 km. S, 3 km. E Rivas, KU 85416; +16 km. S Rivas, MCZ 29009-10; 7.7 km. NE San Juan del Sur, KU 85426-7; +16.5 km. NE San Juan del Sur, KU 85424-5, 87705 (young); 5 km. SE San +Pablo, KU 43079-83. =Zelaya=: Cooley, AMNM 7063-8, 8019-20, 8022, +8034-5; Cukra, AMNH 8016-7; Musahuas, Rio Huaspuc, AMNH 58428-31; 11 km. +NW Rama, Rio Siquia, UMMZ 79708, 79709 (5), 79710 (2); Rio Escondido, +USNM 19766, 20701; Rio Siquia at Rio Mico, UMMZ 79707 (10); Sioux +Plantation, AMNH 7058-61, 8023-33. + +COSTA RICA: =Alajuela=: Los Chiles, AMNH 54639; Orotina, MCZ 7960-1; San +Carlos, USNM 29991. =Guanacaste=: La Cruz, USC 8232 (3); 4.3 km. NE La +Cruz, UMMZ 123002; 18.4 km. S La Cruz, USC 8136; 23.5 km. S La Cruz, USC +8094 (4); 3 km. W La Cruz, USC 8233 (4); 2 km. NE Las Canas, KU 64235-7; +Las Huecas, UMMZ 71212-3; Liberia, KU 36787, USC 8161; 11.5 km. N +Liberia, USC 8149; 13 km. N Liberia, USC 8139; 22.4 km. N Liberia, USC +8126; 8 km. NNW Liberia, KU 64238; 8.6 km. ESE Playa del Coco, USC 8137; +21.8 km. ESE Playa del Coco, USC 8138; Rio Piedra, 1.6 km. W Bagaces, +USC 7027; Rio Bebedero, 5 km. S Bebedero, KU 64158; 5 km. NE Tilaran, KU +36782-6. =Heredia=: 13 km. SW Puerto Viejo, KU 64142-6. =Limon=: Batan, +KU 34927; Guacimo, USC 621; Pandora, USC 505 (3); Suretka, KU 36788-9; +Tortugero, UF 7697, 10540-2. =Puntarenas=: Barranca, CNHM 35254-6; 15 +km. WNW Barranca, KU 64155-7, UMMZ 118381; 18 km. WNW Barranca, UMMZ +118382 (4); 4 km. WNW Esparta, KU 64159-96, 68178-82 (skeletons); 19 km. +NW Esparta, KU 64147-54. + + +=Smilisca cyanosticta= (Smith), new combination + + _Hyla phaeota_, Taylor, Univ. Kansas Sci. Bull., 28(5):80, May 15, + 1942. Taylor and Smith, Proc. U. S. Natl. Mus., 95(3185):589, June + 30, 1945. + + _Hyla phaeota cyanosticta_ Smith, Herpetologica, 8:150, Jan. 30, + 1953 [Holotype.--USNM 111147 from Piedras Negras, El Peten, + Guatemala; Hobart M. Smith collector]. Shannon and Werler, Trans. + Kansas Acad. Sci., 58:386, Sept. 24, 1955. Poglayen and Smith, + Herpetologica, 14:11, April 25, 1958. Cochran, Bull. U. S. Natl. + Mus., 220:57, 1961. Smith, Illinois Biol. Mono., 32:25, May, 1964. + + _Smilisca phaeota cyanosticta_, Stuart, Misc. Publ. Mus. Zool. + Univ. Michigan, 122:42, April 2, 1963. Duellman, Univ. Kansas Publ. + Mus. Nat. Hist., 15(5):229, Oct. 4, 1963. + +_Diagnosis._--Size moderately large ([M] 56.0 mm., [F] 70.0 mm.); skull +as long as wide; frontoparietal fontanelle large; narrow supraorbital +flanges having irregular margins anteriorly; large squamosal not in +contact with maxillary; tarsal fold moderately wide, full length of +tarsus; inner metatarsal tubercle moderately large, low, flat, +elliptical; hind limbs relatively long; tibia usually more than 52 per +cent of snout-vent length; labial stripe silvery-white; lips lacking +vertical bars; loreal region pale green; pale bronze-colored stripe from +nostril along edge of eyelid to point above tympanum narrow, bordered +below by narrow dark brown stripe from nostril to eye, and broad dark +brown postorbital mark encompassing tympanum and terminating above +insertion of arm; flanks, dark brown with large pale blue spots; +anterior and posterior surfaces of thighs dark brown with small +pale blue spots on thighs. (Foregoing combination of characters +distinguishing _S. cyanosticta_ from any other species in genus.) + +_Description and Variation._--The largest males are from Piedras Negras, +El Peten, Guatemala, and they average 52.5 mm. in snout-vent length +whereas males from Los Tuxtlas, Veracruz, average 50.6 mm. and those +from northern Oaxaca 50.3 mm. The smallest breeding male has a +snout-vent length of 44.6 mm. The average ratio of tibia length to +snout-vent length is 54.8 per cent in males from Piedras Negras, +and 56.4 and 56.3 per cent in males from Los Tuxtlas and Oaxaca, +respectively. The only other character showing noticeable geographic +variation is the size of the tympanum. The average ratio of the diameter +of the tympanum to the diameter of the eye is 76.3 per cent in males +from Piedras Negras, 71.8 from Oaxaca, and 66.9 from Los Tuxtlas. + +The dorsal ground color of _Smilisca cyanosticta_ is pale green to tan +and the venter is creamy white. The dorsum is variously marked with dark +olive-green or dark brown spots or blotches (Pl. 6B). An interorbital +dark bar usually is present. The most extensive dark area is a V-shaped +mark in the occipital region with the anterior branches not reaching the +eyelids; this mark is continuous, by means of a narrow mid-dorsal mark, +with an inverted V-shaped mark in the sacral region. In many specimens +this dorsal marking is interrupted, resulting in irregular spots. In +some specimens the dorsum is nearly uniform pale green or tan with a few +small, dark spots. The hind limbs are marked by dark transverse bands, +usually three or four each on the thigh and shank, and two or three on +the tarsus. The webbing on the feet is brown. The loreal region is pale +green, bordered above by a narrow, dark brown canthal stripe extending +from the nostril to the orbit, which is bordered above by a narrow, +bronze-colored stripe extending from the nostril along the edge of the +eyelid to a point above the tympanum. The upper lip is white. A broad +dark brown mark extends posteriorly from the orbit and encompasses the +tympanum to a point above the insertion of the forelimb. The flanks are +dark brown with many pale blue, rounded spots, giving the impression of +a pale blue ground color with dark brown mottling enclosing spots. The +anterior and posterior surfaces of the thighs are dark brown with many +small pale blue spots. The inner surfaces of the shank and tarsus are +colored like the posterior surfaces of the thighs. Pale blue spots are +usually present on the proximal segments of the second and third toes. A +distinct white stripe is present on the outer edge of the tarsus and +fifth toe; on the tarsus the white stripe is bordered below by dark +brown. A white stripe also is present on the outer edge of the forearm +and fourth finger. The anal region is dark brown, bordered above by a +narrow transverse white stripe. The throat in breeding males is dark, +grayish brown with white flecks. + +No geographic variation in the dorsal coloration is evident. Specimens +from the eastern part of the range (Piedras Negras and Chinaja, +Guatemala) have bold, dark reticulations on the flanks enclosing large +pale blue or pale green spots, which fade to tan in preservative. +Specimens from Oaxaca and Veracruz characteristically have finer dark +reticulations on the flanks enclosing smaller blue spots; in many of +these specimens the ventrolateral spots are smallest and are white. + +All living adults are easily recognized by the presence of pale, usually +blue, spots on the flanks and thighs. Individuals under cover by day +have a tan dorsum with dark brown markings. A hiding individual at +Chinaja, Alta Verapaz, Guatemala (KU 55936), had a pale tan dorsum when +found; later the dorsal color changed to chocolate brown. A pale green +patch was present below the eye; the spots on the posterior surfaces of +the thighs were pale blue, and those on the flanks were yellowish green. +A calling male obtained 10 kilometers north-northwest of Chinaja (KU +55934) was reddish brown when found at night; later the dorsal color +changed to pale tan. A green patch below the eye was persistent. Usually +these frogs are green at night. The coloration of an adult male (KU +87201) from 11 kilometers north of Vista Hermosa, Oaxaca, Mexico, was +typical: "When calling dorsum pale green; later changed to dull +olive-green. Flanks dark brown with pale blue spots in axilla and groin +and bluish white flecks on mid-flank. Anterior and posterior surfaces of +thighs, inner surfaces of shanks, and median dorsal surfaces of tarsi +dark brown with blue spots. Canthal and postorbital stripes dark +chocolate brown; labial stripe creamy white. Forearm, tarsal, and anal +stripes pale cream-color. Throat dark brown with yellow flecks; belly +and ventral surfaces of limbs creamy buff; webs pinkish tan; iris deep +bronze, brown below pupil." (Duellman, field notes, June 24, 1964.) + +Some individuals have both green and brown coloration in life. An +individual obtained at night on the south slope of Volcan San Martin, +Veracruz, Mexico, had a pale tan dorsum changing peripherally to pale +green. The dorsal markings were dark brown and dark olive-green. + +In contrast to the color-changes noted above, the labial region below +the eye is always pale green, and pale spots are always present on the +flanks and thighs in adults. The iris is invariably golden or bronze +above and darker, usually brown, below. Minute black flecks are present +on the iris, and in some individuals these flecks are so numerous that +the eye appears gray. + +Recently metamorphosed young have pale tan flanks, and the posterior +surfaces of the thighs are orange-yellow; pale spots are absent. A +juvenile (KU 55935) from Chinaja, Alta Verapaz, Guatemala, having a +snout-vent length of 35.0 mm. was pale yellowish tan above with +olive-green markings; the flanks were dark brown with pale blue spots, +and the anterior and posterior surfaces of the thighs were uniform +bright tomato red. A juvenile (UMMZ 121298), 18.6 mm. in snout-vent +length, from the southeast slope of Volcan San Martin, Veracruz, Mexico, +had pale tan flanks lacking blue spots, but had red thighs. Apparently +the ontogenetic changes in coloration proceed as follows: (1) flanks +pale tan and thighs orange-yellow, both lacking spots, (2) flanks pale +tan and thighs red, both lacking spots, (3) flanks dark brown with blue +spots and thighs red, lacking spots, and (4) flanks and thighs dark +brown, both having pale blue spots. + +_Natural History._--_Smilisca cyanosticta_ inhabits humid tropical +forest and cloud forest from the lowlands to elevations of about 1200 +meters in Los Tuxtlas and to about 900 meters in northern Oaxaca. In +these moist environments the frogs apparently are active throughout the +year. Active individuals have been obtained in January, July, and August +in Los Tuxtlas, Veracruz, in June and July in northern Oaxaca, in +February and March at Chinaja, Guatemala, and Taylor and Smith reported +(1945:589) activity in May at Piedras Negras, Guatemala. Calling males +were observed as follows; in a rain barrel 11 kilometers north of Vista +Hermosa, Oaxaca, Mexico, on June 23-28, 1964; in a quiet pool in a +stream 8 kilometers south of Yetla, Oaxaca, Mexico, in July, 1963 (Dale +L. Hoyt, personal communication); in and near springs flowing into a +stream at Dos Amates, Veracruz, Mexico, on August 4, 1959 (Douglas +Robinson, personal communication); and in a water-filled depression in a +log 10 kilometers west-northwest of Chinaja, Guatemala, on March 13, +1960. Taylor and Smith (1945:589) reported that individuals were found +at night on the ground at the edge of temporary pools at Piedras Negras, +Guatemala, on May 28-29, 1939. A clasping pair was found on a rock at +the edge of a small stream on the south slope of Volcan San Martin, +Veracruz, Mexico, on July 11, 1959 (Douglas Robinson, personal +communication). + +Only one individual has been observed in a tree at night. In the +daytime, individuals were found in elephant ear plants (_Xanthosoma_) at +Chinaja, Guatemala. + +The breeding call consists of one or two moderately short notes that are +lower pitched than those of _S. baudini_, but higher pitched than those +of _S. phaeota_. Each note has a duration of 0.25 to 0.45 seconds and is +repeated at intervals of one-half minute to several minutes. Each note +is a vibrant "waunk," having 110 to 180 pulses per second and dominant +frequency of 1600 to 2100 cycles per second (Pl. 10B). + +Apparently the eggs are deposited as loose clumps in the water. In eggs +in the yolk plug stage of development, the diameter of the embryo is +about 2.3 mm.; that of the outer envelope is 4.0 mm. Hatchling tadpoles +have total lengths of 5.8 to 6.5 mm. and body lengths of 2.8 to 3.1 mm. +The external gills are moderately long, slender, and filamentous; the +yolk sac is still moderately large. The body and anterior part of the +caudal musculature are dark brown; posteriorly the caudal musculature is +pale brown. The caudal fins are creamy tan. The oral discs are large and +ovoid. The growth of the tadpole is summarized in Table 10. + +A typical tadpole in stage 30 of development (KU 87652 from 11 km. N +Vista Hermosa, Oaxaca, Mexico) can be described as follows: + +Body length 9.5 mm.; tail length 15.5 mm.; total length 25.0 mm.; body +slightly wider than deep; snout rounded laterally, broadly ovoid +dorsally; eyes widely separated, directed dorsolaterally; nostril about +midway between eye and tip of snout; mouth anteroventral; spiracle +sinistral, slightly posterior to midpoint of body and slightly below +midline; anal tube dextral; caudal musculature slender, barely curved +upward distally; dorsal fin not extending onto body, depth of dorsal fin +slightly more than that of ventral fin on mid-length of tail; dorsal +part of body dark brown; ventral surfaces transparent, lacking pigment; +posterior edge of body pale cream-color; caudal musculature creamy white +with interconnected brown spots; caudal fins transparent with small +brown blotches on dorsal fin and posterior half of ventral fin; iris +coppery bronze in life (Fig. 12). Mouth small, median part of upper lip +bare; rest of mouth bordered by single row of bluntly rounded papillae; +lateral fold present; tooth rows 2/3; all tooth-rows approximately equal +in length; second upper row broadly interrupted medially; other rows +complete; upper beak moderately deep, forming broad arch with slender +lateral processes; lower beak slender, broadly V-shaped; both beaks +finely serrate (Fig. 15C). + +All tadpoles having fully developed mouth parts have 2/3 tooth rows. +Little variation is noticeable in coloration. In many specimens the +posterior edge of the body is dark brown instead of pale cream-color. +Mottling is rather dense on the caudal fins in all specimens; in some +individuals pigment is concentrated along the anterior one-third of the +lateral groove. In life the body is dark brown with greenish gold flecks +ventrally; the caudal musculature is gray. + +In each of two recently metamorphosed young the snout-vent length is +14.0 mm. Coloration of young in life (KU 87653 from 11 km. N Vista +Hermosa, Oaxaca, Mexico): "Dorsum pale tan with dark brown markings. +Thighs orange-yellow; labial stripe white; iris bronze" (Duellman, field +notes, July 10, 1964.) + +_Remarks._--Smith (1953:150) named _cyanosticta_ as a subspecies of +_Hyla phaeota_. The differences in cranial characters and certain +external characters between _phaeota_ and _cyanosticta_ indicate that +they are distinct species. Furthermore, a gap of about 350 kilometers +exists between the known geographic ranges of the two kinds. + +_Distribution._--_Smilisca cyanosticta_ inhabits wet forests on the +Atlantic slope of southern Mexico and northern Central America from +northern Oaxaca and southern Veracruz through northern Chiapas in Mexico +and into El Peten and northern Alta Verapaz in Guatemala (Fig. 2). +Apparently the range is discontinuous, for in southern Mexico the +species is found in cloud forest at elevations of 830 to 900 meters on +the northern slopes of the Sierra de Juarez. In the Sierra de Los +Tuxtlas in southern Veracruz the species is found in wet forest at +elevations of 300 to 1200 meters, but is absent in the intervening +lowlands characterized by drier forest. In the west forests of northern +Alta Verapaz and El Peten, Guatemala, the species is found at low +elevations. + +_Specimens examined._--78, as follows: MEXICO: =Chiapas=: Monte Libano, +MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH +41588. =Oaxaca=: 11 km. N Vista Hermosa, KU 84918-20 (skeletons), +87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH +57199-201; 8 km. S Yetla, KU 87213, UMMZ 124838 (8). =Veracruz=: Coyame, +UMMZ 111459-60; between Coyame and Tebanco, UMMZ 121198; Dos Amates, +UMMZ 121297; between Laguna de Catemaco and Volcan San Martin, UMMZ +121200; Volcan San Martin, UIMNH 35403-4, 35408-12, UMMZ 118163; SE +slope Volcan San Martin, UMMZ 121199, 121295 (2), 121296, 121298. + +GUATEMALA: =Alta Verapaz=: Chinaja, KU 55935-7, 55938 (skeleton). =El= + +=Peten=: 10 km. NNW Chinaja (Alta Verapaz), KU 55934; Piedras Negras, +CNHM 99006-7, 99227, UIMNH 28853, USNM 111139-41, 111143-7; 8 km. S +Piedras Negras, CNHM 99008; Semicoch, USNM 35907. + + [Illustration: 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. + Muenchen, 27:215, 1897. Guenther, Biologia Centrali-Americana: + Reptilia and Batrachia, p. 269, Sept. 1901. Nieden, Das Tierreich, + Amphibia, Anura I, p. 261, June 1923. Dunn, Occas. Papers Boston + Soc. Nat. Hist., 5:413, Oct. 10, 1931. Gaige, Hartweg, and Stuart, + Occas. Papers Mus. Zool. Univ. Michigan, 357:4, Oct. 26, 1937. + Cooper, Copeia, 2:122, June 30, 1944. Breder, Bull. Amer. Mus. Nat. + Hist., 86(6):416, Aug. 26, 1946. Smith and Taylor, Bull. U. S. Natl. + Mus., 194:88, June 17, 1948; Univ. Kansas Sci. Bull, 33:364, March + 20, 1950. Taylor, Univ. Kansas Sci. Bull., 35(1):837, July 1, 1952. + Brattstrom and Howell, Herpetologica, 10:117, Aug. 1, 1954. Goin, + Herpetologica, 14:120, July 23, 1958. Cochran, Bull. U. S. Natl. + Mus., 220:57, 1961. + + _Hyla labialis_ Peters, Monats. Konigl. Akad. Wissen. Berlin, p. + 463, 1863 [Holotype.--ZMB 4913 from "region of Bogota," Colombia]; + Monats. Konigl. Akad. Wissen. Berlin, p. 618, Oct. 16, 1873. + Boulenger, Catalogue Batrachia and Salientia in British Museum, p. + 397, Feb. 1, 1882. + + _Hyla baudini dolomedes_ Barbour, Occas. Papers Mus. Zool. Univ. + Michigan, 129:11, Jan. 25, 1923 [Holotype.--MCZ 8539 from Rio + Esnape, Sambu Valley, Darien, Panama; Barbour and Brooks + collectors]. Barbour and Loveridge, Bull. Mus. Comp. Zool. Harvard, + 69:278, June, 1929. + + _Hyla phaeota phaeota_, Smith, Herpetologica, 8:152, Jan. 30, 1953. + Minton and Smith, Herpetologica, 16:103, June 17, 1960. + + _Smilisca phaeota_, Starrett, Copeia, 4:303, Dec. 30, 1960. + +_Diagnosis._--Size large ([M] 65 mm., [F] 78 mm.); skull as long as +wide, lacking frontoparietal fontanelle; large supraorbital flanges +having straight edges and extending posterolaterally; large squamosal +not in contact with maxillary; tarsal fold moderately wide, full length +of tarsus; inner metatarsal tubercle moderately large, low, flat, +elliptical; hind limbs relatively long, tibia averaging more than 53 per +cent of snout-vent length; labial stripe silvery white; lips lacking +vertical bars; loreal region pale green; dark brown or black tympanic +mark dispersing into brown venated pattern on flanks; posterior surfaces +of thighs pale brown, with or without darker flecks or small +cream-colored flecks. (Foregoing combination of characters +distinguishing _S. phaeota_ from any other species in genus.) + + + Table 2.--Geographic Variation in Size and Proportions in Males of + Smilisca phaeota. (Means in Parentheses Below Observed Ranges; + Data Based on Ten Specimens From Each Locality.) + + ================================================================== + |Snout-vent |Head width/|Interorbital + Locality |length |snout-vent |distance/ + | |length |head width + -----------------------------+-----------+-----------+------------ + Bonanza, Nicaragua | 40.8-47.7 | 34.1-38.0 | 31.0-36.1 + | (43.7) | (36.3) | (35.4) + -----------------------------+-----------+-----------+------------ + Heredia Prov., Costa Rica | 46.3-59.0 | 32.5-36.0 | 30.5-39.6 + | (51.7) | (35.0) | (34.7) + -----------------------------+-----------+-----------+------------ + Puntarenas Prov., Costa Rica | 53.6-64.9 | 32.6-36.1 | 31.0-38.0 + | (61.4) | (34.5) | (34.4) + -----------------------------+-----------+-----------+------------ + Canal Zone, Panama | 52.4-65.5 | 33.5-37.6 | 31.3-37.2 + | (56.5) | (35.6) | (34.7) + -----------------------------+-----------+-----------+------------ + Rio Quesada, Colombia | 52.6-61.0 | 33.1-37.1 | 30.1-33.9 + | (56.0) | (35.0) | (32.1) + ------------------------------------------------------------------ + + +_Description and Variation._--For the purposes of analyzing geographic +variation in size and proportions, measurements were taken on ten adult +males from each of five samples throughout the range of the species. +Aside from the data summarized in Table 2, the average ratio of tibia +length to snout-vent length is noticeably less in Colombian specimens +(53.4 per cent, as compared with 54.8 to 57.8 per cent in the other +samples) and the ratio of head length to snout-vent length is +noticeably less in Costa Rican specimens (33.5 per cent as compared with +34.9 to 35.1 per cent in the other samples). Also, specimens from +Heredia Province, Costa Rica, have a relatively smaller tympanum (62.7 +to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared +with means of 74.0 to 77.9 per cent in the other samples). + +Two populations are distinctive as regards the size of adult males. +Specimens from the northern Caribbean lowlands of Nicaragua (Bonanza, +the northernmost locality for the species) are remarkably small. Males +having snout-vent lengths of between 40 and 43 mm. were breeding; the +largest male found had a snout-vent length of 47.7 mm. The other extreme +in size is attained in specimens from the Pacific lowlands of eastern +Costa Rica and western Panama, where most breeding males have snout-vent +lengths of more than 55 mm.; the largest male had a snout-vent length of +64.9 mm. + +The rather striking differences in size among certain samples and the +minor differences in proportions among other samples show no geographic +trends. Instead, the variations apparently are random among the samples. +The data presented here possibly are the results of inadequate sampling, +but more likely reflect actual differences in the populations. + +The dorsal ground color of _Smilisca phaeota_ is pale green to tan; the +venter is creamy white. The dorsum is variously marked with dark +olive-green or dark brown spots or blotches (Pl. 6C). A dark +interorbital bar is usually present. Usually a large dark dorsal mark +extends from the occiput to the sacral region, but in many individuals +this blotch is replaced by two or three dark marks. The dorsal markings +are irregular in shape and do not tend to form transverse bands or +longitudinal bars. The hind limbs are marked by dark transverse bands, +usually four or five on the thigh, five or six on the shank, and four on +the tarsus. Two or three narrow bands are usually present on the +proximal part of the fourth toe. The webbing on the feet is brown. The +loreal region is pale green, bordered above by a narrow dark brown +canthal stripe extending from the nostril to the orbit. The upper lip is +silvery white. A broad dark brown or black mark extends posteriorly from +the orbit, encompassing the tympanum, to a point above the insertion of +the forelimb. The flanks are pale green or pale tan and marked with a +fine dark brown or black venation. The anterior surfaces of the thighs +usually are pale brown or grayish tan, sometimes having small, +indistinct darker flecks. The posterior surfaces of the thighs are +similarly colored, but in most specimens small but distinct dark flecks +are present; in some specimens small cream-colored spots are also +present on the posterior surfaces of the thighs. A distinct, narrow +creamy white anal stripe usually is present. A distinct white stripe is +present on the outer edge of the tarsus and fifth toe; on the tarsus the +white stripe is bordered below by dark brown. A white stripe also is +present on the outer edge of the forearm and fourth finger. In breeding +males the throat is dark gray. + +Little geographic variation in color or pattern is evident. Few, if any, +specimens from the Pacific lowlands of South America are green in life. +(We have seen no living individuals from South America.) Some living +individuals from Costa Rica and all those seen alive from Nicaragua have +a tint of pale blue on the flanks. In some specimens the dorsal pattern +is so faint as to be barely discernible, whereas in most specimens the +pattern is bold. + +The coloration in the living frogs is highly variable due to extreme +metachrosis. Individuals of this species are capable of changing the +dorsal coloration from green to brown in a short period of time. Both +green and brown individuals have been found active at night. Usually +those individuals found hiding by day are brown. One individual from +Finca La Sumbadora, Panama (now KU 91914), was kept alive in the +laboratory for nearly one month. This individual usually was pale green +with tan dorsal markings at night and tan with pale green markings by +day. On occasion the pale green dorsal markings were boldly outlined by +bright dark green. + +In living individuals from throughout the range of the species the iris +is a bronze color, darkest medially with fine black reticulations. + +_Natural History._--_Smilisca phaeota_ inhabits humid lowland tropical +forest and seldom ascends the foothills to more than 1,000 meters. The +rather equable climatic conditions, especially more or less evenly +distributed rainfall throughout the year, permit this frog to be active +most of the year. Dunn (1931:413) reported males calling on Barro +Colorado Island, Panama, in February and in July, and Breder (1946:416) +noted calling individuals in the Chucanaque drainage of Darien, Panama +in January, March, July, August and October and in Costa Rica in April +through August inclusively. Calling males were found at Bonanza, +Nicaragua in March and in July. + +At all times of year the usual daytime retreats for these frogs are near +water; the frogs have been found in elephant ear plants (_Xanthosoma_) +and in bromeliads; occasional individuals have been found sitting on +shaded branches of bushes and trees. None has been observed on the +ground or beneath ground-cover by day. + +The length of the breeding season cannot be determined definitely. The +earliest date on which eggs have been found is May 23; Gaige, Hartweg, +and Stuart (1937:5) reported a gravid female taken at El Recreo, +Nicaragua, in September, and we have a gravid female taken at Almirante, +Panama, in March. + +Males usually call from secluded spots at the edge of water. All calling +males that we observed were on the ground within a few centimeters of +the water. The males usually are hidden beneath an overhanging leaf or +some other cover; they definitely do not sit in the open like _Smilisca +baudini_. Most calling males and clasping pairs have been found at the +edges of small pools or shallow ditches, although occasional individuals +are found at the edges of large ponds or streams. + +The breeding call consists of one or two moderately short, low-pitched +notes (duration 0.33 to 0.42 seconds), repeated at intervals of about 20 +seconds to several minutes. Each note is a low, vibrant "wauk," having +100 to 130 pulses per second and a dominant frequency of 330 to 420 +cycles per second (Pl. 10C). + +The eggs are deposited in loose clumps amidst vegetation in the water. +Hatchling tadpoles have total lengths of 8.7 to 10.6 mm., and body +lengths of 4.1 to 4.5 mm. The external gills are long and filamentous, +and the yolk sac is large. The head and caudal musculature are dark +brownish black, and the caudal fins are gray. The oral discs are large +and roughly circular. The growth and development of the tadpoles are +summarized in table 11 and figure 16. + +A typical tadpole in stage 30 of development (KU 68482 from the Rio +Chitaria, Cartago Province, Costa Rica) may be described as follows: +body length 9.7 mm.; tail length 14.6 mm.; total length 24.3 mm.; body +as wide as deep; snout rounded dorsally and laterally; eyes widely +separated, directed dorsolaterally; nostril about midway between eye +and tip of snout; mouth anteroventral; spiracle sinistral, about midway +on length of body and slightly below midline; anal tube dextral; caudal +musculature slender, curved upward distally; dorsal fin extending onto +body; depth of dorsal fin slightly less than that of ventral fin at +mid-length of tail; dorsal part of body pale brown; ventral surfaces +transparent with scattered pigment; pale cream-colored, crescent-shaped +mark on posterior edge of body; caudal musculature pale creamy tan with +scattered pale brown spots; caudal fins transparent with scattered small +brown blotches on dorsal and ventral fins; iris pale bronze in life +(Fig. 13); mouth small; median part of upper lip bare; rest of mouth +bordered by one row of pointed papillae; lateral fold present; +tooth-rows 2/3, first upper row longest; second upper row slightly +shorter, broadly interrupted medially; three lower rows complete, equal +in length, slightly shorter than second upper row; upper beak moderately +deep, forming broad arch with slender lateral processes; lower beak +slender, broadly V-shaped; both beaks serrate (Fig. 15E). + +In tadpoles having fully developed mouthparts the tooth-row formula of +2/3 is invariable. The pale crescent-shaped mark on the posterior part +of the body curves anterodorsally on the dorsal surface of the body. +These marks in dorsal view give the appearance of a pair of short, pale +stripes on the posterior part of the body. Most specimens from Costa +Rica have the pale coloration like that described above, but some +individuals (notable KU 87683 from Guapiles, Costa Rica, KU 87707 from +Finca Tepeyac, Nicaragua, and KU 87708 from Bonanza, Nicaragua) have +much more pigment. In these specimens the same color pattern obtains as +in the pallid individuals, but the pigmentation is dense. This is +especially noticeable on the tail. + +Recently metamorphosed young have snout-vent lengths of 12.7 to 16.7 mm. +(average, 14.3 mm. in eleven specimens). Coloration of young in life (KU +68484 from Rio Chitaria, Cartago Province, Costa Rica): "Dorsum pale +tan; side of head and flanks darker brown, separated from tan dorsum by +an indistinct cream stripe. Limbs pale yellow; thighs flecked with +brown; shank and tarsus yellowish tan with indistinct brown bars. Soles +of feet brown. Belly white; throat dusty cream flecked with silvery +white. Upper lip silvery white. Iris bright gold with black flecks. +Heels, tarsal and anal stripes white" (Duellman, field notes, May 23, +1961). + +_Remarks._--Peters (1863:463) named _Hyla labialis_ from the "region of +Bogota, Colombia", but in 1873 regarded his new species as identical +with _Hyla phaeota_ Cope, 1862, from Turbo, Colombia. The name _Hyla +labialis_ has been used for frogs from the northern Andes in Colombia +(see Dunn, 1944:72, and Stebbins and Hendrickson, 1959:522, for +discussion of nomenclature). Rivero (1961:131) used the name _Hyla +vilsoniana_ Cope, 1899, for the frogs from the northern Andes previously +referred to _Hyla labialis_. A review of the nomenclature and taxonomy +of these frogs, which superficially resemble _Smilisca_ but are +unrelated, is beyond the scope of the present study. + +_Hyla baudini dolomedes_ Barbour, 1923, is based on a small _Smilisca +phaeota_ (MCZ 8539) having a snout-vent length of 45.5 mm. Dunn +(1931a:413) placed _dolomedes_ in the synonymy of _Smilisca phaeota_. We +have examined the holotype of _dolomedes_ and agree with Dunn's +assignment. + +Smith (1953:150) described _Hyla phaeota cyanosticta_ from Guatemala. +Our studies on the external morphology, coloration, and especially the +cranial osteology provide evidence that _cyanosticta_ is a species +distinct from _phaeota_. + +_Distribution._--_Smilisca phaeota_ inhabits humid tropical forests from +northeastern Nicaragua southward on the Caribbean lowlands to elevations +of about 1000 meters and on the Pacific lowlands of Costa Rica, +exclusive of the arid regions of Guanacaste, throughout the lowlands of +Panama, exclusive of the savannas of the Pacific lowland and the Azuero +Peninsula, and southward on the Pacific slopes of South America through +Colombia to west-central Ecuador; also the valleys of the Rio Cauca and +Rio Magdalena in Colombia (Fig. 2). + +_Specimens examined._--528, as follows: NICARAGUA: =Matagalpa=: Finca +Tepeyac, 10 km. N, 9 km. E Matagalpa, KU 85439, 87707 (tadpoles); +Matagalpa, MCZ 3546-7, UMMZ 92367; 19 km. N Matagalpa, UMMZ 116495-6. +Zelaya: Bonanza, KU 84854-62, 84950-2 (skeletons), 85440-50, 87708-9 +(tadpoles); Cukra, AMNH 80618; Rio Mico, 16 km. E Recreo, UMMZ 79711 +(6), 79712 (4); junction Rio Mico and Rio Siguia, UMMZ 79713 (10); Rio +Siguia, 11 km. NW Rama, UMMZ 79714 (14), 79715 (11), 79716 (21), 79717, +79718 (3). + +COSTA RICA: =Alajuela=: Cinchona, KU 32255, 64286-8; 5 km. S Ciudad +Quesada, USC 8077; Laguna Monte Alegre, KU 64289-90; Las Playuelas, 11 +km. S Los Chiles, USC 7216; San Carlos, USNM 29961. + +=Cartago=: Moravia de Turrialba, KU 32212-47, 37133-5, 41093 (skeleton), +64280-1, USC 7243 (3); Peralta, KU 32271-2; Rio Chitaria, 3 km. NNE +Pavones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); +Rio Reventazon, MCZ 29196-203, UMMZ 117677 (9); Turrialba, KU 25720-2, +32209-11, 32266-8, 32273-4, 37136-67, 41090-2 (skeletons), 64270-2, MCZ +29221, 29222 (tadpoles), 29269-70, USNM 29934. + +=Guanacaste=: Tilaran, KU 36805-7; 8 km. NE Tilaran, KU 36803-4. + +=Heredia=: Barranca del Rio Sarapiqui below Isla Bonita, KU 64282-3; +Cariblanco, KU 32256-60, 41094 (skeleton), 64284, MCZ 7967; Isla Bonita, +KU 32250-4; 4.2 km. W Puerto Viejo, KU 64285, 68485; 7.5 km. W Puerto +Viejo, KU 68486; 1 km. S Puerto Viejo, KU 86518. + +=Limon=: Bambu, USC 7182 (4); Batan, UMMZ 118582; Coen, MCZ 9825; La +Lola, KU 32262-4, UF 4029, UMMZ 117678 (3); Los Diamantes, CNHM +101295-8, KU 25723-4, 32265, 64267-9; Pandora, UMMZ 122650 (2), USC 7188 +(3), 7190; Puerto Limon, KU 32261; Rio Lari at Rio Dipari, 21 km. SW +Amubre, USC 7177; Rio Toro Amarillo, 7 km. W Guapiles, KU 86519, 87683 +(tadpoles); Suretka, KU 36808-10, 37168. + +=Puntarenas=: Agua Buena, KU 36790; 1.6 km. E Buenos Aires, UMMZ 117578; +3 km. NW Buenos Aires, KU 64304; 4 km. N, 15 km. W Dominical, KU 68491-2 +(tadpoles); Esparta, MCZ 8029-30, 8032; Golfito, KU 32270; 6 km. E +Golfito, KU 84999-500 (skeletons); Gromaco, UMMZ 123677 (4); Palmar, KU +32269; 4 km. ESE Palmar Sur, KU 64305-6; 5.6 km. SE Palmar Sur, KU 68489 +(tadpoles); 7.0 km. SE Palmar Sur, KU 68490 (young); 8.5 km. SE Piedras +Blancas, KU 64292-303; Quebrada Boruca, 22 km. E Palmar Norte, KU 64291; +Rincon, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), USC 7254; Rio +Ferruviosa, 7 km. S Rincon, USC 7235; 1.6 km. WNW Villa Neily, KU 68493 +(young), 68494 (tadpoles). + +=San Jose=: San Isidro el General, KU 32249, UMMZ 75025; 10 km. N San +Isidro el General, MCZ 29099-103; 13 km. WSW San Isidro el General, KU +86517; 15 km. WSW San Isidro el General, KU 68487 (tadpoles), 68488 +(young), 68495 (young); 20 km. WSW San Isidro el General, KU 32248. + +PANAMA: No province: Cano Saddle, USNM 69588; Punta de Pena, USNM 38733; +Quipo, AMNH 18925-6. =Bocas del Toro=: Almirante, KU 80080, 91835-6; 1.6 +km. W Almirante, KU 91837; 3 km. W Almirante, KU 91824 (skeleton), +91838-43, 91906-7; 11 km. NW Almirante, CNHM 67853-61; 13 km. W +Almirante, KU 91825-7 (skeletons), 91844-9; Fish Creek, KU 92329; Isla +Popa, KU 91850-1. =Canal Zone=: Barro Colorado Island, CNHM 6007, 13316, +13325, 13331, 13360-2, 13377-8, MCZ 24191-5, UF 7523, UMMZ 63547-60, +64457, 69497 (3); 3.7 km. W Cocoli, KU 67916; Fort Sherman, MCZ 10139; +Gatun, MCZ 35644; Junction roads C25B and C16, TNHC 23839; Madden +Forest Preserve, TNHC 23837-8. =Cocle=: El Valle, KU 77521-4, 77649 +(tadpole), TNHC 23369. =Comarca del Baru=: Progreso, UMMZ 61085-9. +Colon: Achiote, KU 77516-20, 77648 (young); Rio Candelaria, CNHM +67851-2. =Darien=: Rio Esnape, Sambu Valley, MCZ 8539; Rio Sucubti, +Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, +Camp Townsend, AMNH 40988. =Panama=: NW slope Cerro Prominente, KU +80459; Finca La Sumbadora, KU 91914 (skeleton). =Chiriqui=: 2 km. W +Concepcion, AMNH 68910. + +COLUMBIA: =Antioquia=: Puerto Berrio, CNHM 30805 (Goin); Turbo, USNM +39899. =Caldas=: Pueblorrica, Santa Cecilia, CNHM 54768-71 (Goin). +=Choco=: No specific locality, AMNH 3984-6; Andagoya, BMNH 1915.10.21. +69-70, CNHM 81857 (Goin); Golfo de Uraba, CNHM 63881 (Goin); Pena Lisa, +Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro, +CNHM 4451-3, 4455-61 (Goin); Rio San Juan, Playa del Oro, CNHM 54772 +(Goin); Rio Quesada, AMNH 13615-77; 37 km. up Rio Pune, AMNH 13688; 48 +km. up Rio Pune, AMNH 13689. =Narino=: Tumaco, Rio Rosario, CJG 2310-13 +(Goin). =Valle=: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT +166, 346-47, 388 (Goin); Rio Calima above Cordoba, CJG 2249-57 (Goin). + +ECUADOR: No province: Bulun, AMNH 10620. =Esmeraldas=: Cachabe, AMNH +10625-8; Rio Capayas, CNHM 35712; Rio Sapaya, UMMZ 58910 (5); Salidero, +AMNH 10623-4; San Javier, AMNH 10618. =Guayas=: Hacienda Balao Chico, +UMMZ 123904. =Imbabura=: Pambelar, AMNH 10629, 10631. =Pichincha=: +Hacienda Espinosa, 9 km. W Santo Domingo de los Colorados, KU 40220. + + +=Smilisca puma= (Cope), new combination + + _Hyla puma_ Cope, Proc. Amer. Philos. Soc., 22:183, 1885 + [Holotype.--USNM 13735 from Nicaragua; J. F. Moser collector]. + Guenther, Biologia Centrali-Americana: Reptilia and Batrachia, p. + 270, Sept., 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 251, + June, 1923. Cochran, Bull. U. S. Natl. Mus., 220:58, 1961. + + _Hyla wellmanorum_ Taylor, Univ. Kansas Sci. Bull. 25(1):843, July + 1, 1952 [Holotype.--KU 30302 from Batan, Limon, Costa Rica, Edward + H. Taylor collector]; Univ. Kansas Sci. Bull., 36(1):626, June 1, + 1954. Duellman and Berg, Univ. Kansas Publ. Mus. Nat. Hist., + 15:194, Oct. 26, 1962. + + _Smilisca wellmanorum_, Starrett, Copeia, 4:303, Dec. 30, 1960. + +_Diagnosis._--Size small ([M] 38.0 mm., [F] 46.0 mm.), differing from +other species in the genus by the following combination of characters: +skull about as long as broad; frontoparietal fontanelle keyhole-shaped; +supraorbital flanges absent; squamosal small, not in contact with +maxillary; bony portion of ethmoid terminating at anterior edge of +orbit; tarsal fold weak, two-thirds length of tarsus; inner metatarsal +tubercle small, low, flat, elliptical; snout rounded in dorsal profile; +lips thin and flaring; fingers having only vestige of web; toes one-half +webbed; diameter of tympanum about two-thirds that of eye; narrow labial +stripe white; pair of dark brown (sometimes interconnected) stripes on +tan dorsum; no blue spots on flanks or thighs; vocal sac in breeding +males pale brown. (Foregoing combination of characters distinguishing +_S. puma_ from other species in genus.) + +_Description and variation._--Ten breeding males from the vicinity of +Puerto Viejo, Heredia Province, Costa Rica, have snout-vent lengths of +32.5 to 37.9 mm. (34.8 mm.). In these specimens, the length of the tibia +to the snout-vent length is 0.48 to 0.53 (0.51), and the tympanum/eye +ratio is 0.52 to 0.72 (0.65). Seven females have snout-vent lengths of +40.8 to 45.8 mm. (43.9 mm.). No individual has more than a vestige of a +web between the second and third and fourth fingers. None has a web +between the first and second fingers. Breeding males lack nuptial +excrescences on the thumbs. The vocal sac is moderately large and +bilobed. + +In preserved specimens the dorsal ground color varies from yellowish tan +to grayish brown. All specimens have dark brown dorsal markings in the +form of a pair of dorsal stripes, variously modified (Pl. 7A). In some +specimens, such as KU 91716, the stripes are discrete and extend from +the postorbital region nearly to the vent. In most specimens the stripes +are connected by a transverse mark in the scapular region and in many +others also by a transverse mark in the sacral region. In some specimens +the stripes are fragmented posteriorly; fragmentation is extreme in KU +30300, in which the dorsal pattern consists of two series of dark +longitudinal dashes. The other extreme is a nearly complete fusion of +the stripes, as in KU 91714. A dark brown interorbital bar usually +extends onto the eyelids, but in some specimens this is reduced to a +short V-shaped mark or small spot between the eyes. There is no dark +post-tympanic mark, but dark brown pigment forms a venated pattern from +the axilla to the mid-flank; the inguinal region is white, finely +mottled with dark brown. The dorsal surfaces of the hind limbs are +colored like the body and have two or three dark brown transverse marks +on the thighs, three to five marks on the shanks, and one or two marks +or irregularly arranged dark flecks on the tarsi. The anterior and +posterior surfaces of the thighs are pale tan to brown. The webbing of +the feet is tan to grayish brown. A narrow white labial stripe, white +anal stripe, and narrow white stripes on the tarsi and outer edges of +the forelimbs are invariably present. The ventral surfaces are creamy +white. + +In life the dorsum is tan or pale brown with dark brown markings. Some +individuals have scattered metallic green flecks on the dorsum. The +flanks are mottled dark brown and creamy white. The posterior surfaces +of the thighs are dark brown. The vocal sacs are grayish brown, and the +iris is a deep bronze color. + +_Natural History._--_Smilisca puma_ inhabits humid lowland tropical +forests having more or less evenly distributed rainfall throughout the +year. The equable climatic conditions seemingly permit these frogs to be +active throughout most of the year. Taylor (1952:846) found calling +males at Batan, Costa Rica, on July 20, 1951. We found the species +breeding near Puerto Viejo, Costa Rica, on February 19, June 18, July +13, and July 31. Specimens of calling males from Costa Rica in the +collection at the University of Southern California were obtained in +February at La Fortuna, on August 22 at Los Diamantes, on August 30 at +Jabillos, and on September 5 at La Lola. Gravid females were collected +in June, July and August. + +Males call from shallow water. All breeding congregations of this +species that we have found were in a grassy marsh, 7.5 kilometers west +of Puerto Viejo, Costa Rica. Tadpoles were found in water-filled +depressions in the marsh at night. When first observed, tadpoles were +near the surface of the water; they responded to light by quickly taking +refuge in the dense grass. No tadpoles were observed by day. + +The breeding call consists of a low squawk, usually followed by a series +of one or more rattling secondary notes (duration of primary notes, +0.06-0.35 seconds; of secondary notes, 0.10 to 0.47 seconds), repeated +at intervals of 5 to 55 seconds. The primary notes have 187 to 240 +pulses per second and major frequencies of about 740 to 1870 cycles per +second (Pl. 11A). + +Only six tadpoles are available for study. Four of them in stage 34 of +development have body lengths of 9.0 to 9.5 mm., tail lengths of 14.0 to +15.0 mm., and total lengths of 23.0 to 24.5 mm. One tadpole in stage 38 +and one in stage 40 have total lengths of 31.0 mm. A typical tadpole in +stage 34 of development (KU 91807 from 7.5 km. W Puerto Viejo, Heredia +Province, Costa Rica) has a body length of 9.5 mm., tail length of 15.0 +mm., and total length of 24.5 mm.; body about three-fourths as deep as +wide; snout rounded dorsally and laterally; eyes widely separated, +directed dorsolaterally; nostril about midway between eye and tip of +snout; mouth anteroventral; spiracle sinistral, about two-thirds +distance from snout to posterior end of body and slightly below midline; +anal tube dextral; caudal musculature slender, barely curved upward +distally; dorsal fin extending onto body; at mid-length of tail, depth +of caudal musculature equal to that of dorsal fin and ventral fin; body +grayish brown, palest ventrally; caudal musculature pale creamy yellow +with bold gray reticulations; caudal fins transparent with gray +reticulations anteriorly and black flecks posteriorly on both fins (Fig. +14A). Median part of upper lip bare; rest of mouth bordered by two rows +of short blunt papillae; lateral fold present; tooth-rows 2/3; upper +rows equal in length; second upper row broadly interrupted medially; +three lower rows complete, first and second rows equal in length, +slightly shorter than upper rows; third lower row noticeably shorter; +upper beak shallow, forming broad, continuous arch with slender lateral +processes; lower beak slender, broadly V-shaped, both beaks finely +serrate (Fig. 15B). + +All six tadpoles are colored alike, except that in the larger specimens +scattered white flecks are present on the ventral surface of the body, +and the dark reticulations continue farther posteriorly on the caudal +fins than in the smaller tadpoles. In two specimens the third lower +tooth-row is only about one-half the length of the other lower rows, and +in one specimen the second lower tooth-row is shorter than the first. +Coloration of tadpoles in life: "Body olive-brown with silvery green +flecks laterally. Caudal musculature olive-brown with greenish tan +flecks. Fins brown with greenish gold flecks. Iris deep bronze." +(Duellman, field notes, February 19, 1965). + +One recently metamorphosed young (KU 91808) has a snout-vent length of +12.4 mm. In life this frog had a pale tan dorsum with dark brown +markings, yellowish tan posterior surfaces of thighs, grayish brown +throat, and bronze iris. + +_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. + + [Illustration: 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); Rio 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=: Batan, 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 Volcan, Chiriqui Province, Panama, 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 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, Panama, 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 Panama). 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 Volcan, Chiriqui, Panama, 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 Volcan +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 +Panama. 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 Panama 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 +Panama. In specimens from Costa Rica the flanks are brown with pale blue +flecks, whereas in those from Chiriqui, Panama, 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 Chiriqui, 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 Darien and +from Panama Province east of the Canal Zone (Altos de Pacora, Cerro +Jefe, Finca La Sumbadora, and Rio 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, Panama, 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, Panama, 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 Panama west of the Canal Zone, +the blue coloration on the flanks and thighs is much less conspicuous +than in specimens from eastern Panama. 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, Chiriqui, Panama, +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 Volcan, Chiriqui, Panama). 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, Panama) 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 Volcan, Chiriqui (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 +Panama 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 Panama 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; +Rio Zapote, 7 km. E Palmar Norte, USC 7100 (2). =San Jose=: 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. =Chiriqui=: Boquete, AMNH 69815, +UMMZ 58441-5; El Volcan, KU 77413, 91828-31 (skeletons), 91852-74, 91832 +(eggs), 91833 (tadpoles); 6 km. S El Volcan, CNHM 60442; 16 km. NNW El +Volcan, KU 91879-90; Finca Palosanto, 6 km. WNW El Volcan, KU 77406-12, +77692 (skeleton), 91875-7, 92330-1; Rio Colorado, 17 km. NNW El Volcan, +KU 91878, 92332; Valle Hornito, 19 km. NE Gualaca, KU 92333-45. =Cocle=: +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. =Colon=: Rio Candelaria, AMNH 53708-15, CNHM 67826-36. =Darien=: +Camp Creek, Camp Townsend, AMNH 40756-7, 40936-9, 40992; Rio Chico, AMNH +39784, 40986-7; Rio Pita, CNHM 67823-5; Tacarcuna, USNM 141796-802; +Three Falls Creek, AMNH 41684, 51788. =Los Santos=: Cerro Hoya, USNM +148213-4; Lajamina, Rio Puria, KU 67915. =Panama=: 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); Rio Calobra, USNM 53722, Rio Pacora, 9 km. NNE Pacora, KU +91901. =Veraguas=: Cerro Carbunco, USNM 129066; Cerro Tute, CNHM +67837-41; Isla Cebaco, Rio Platanal, KU 91891-3. + +COLOMBIA: =Antioquia=: Uraba, Villa Arteaga, CNHM 63893 (Goin). +=Atlantico=: Sabanalarga, Rio 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," + Panama; J. von Warszewicz collector]. Brocchi, Mission scientifique + au Mexique ..., pt. 3, sec. 2, Etudes sur les batrachiens, p. 42, + 1881. Boulenger, Catalogue Batrachia Salientia in British Museum, + p. 393, Feb. 1, 1882. Guenther, 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, Limon, + Costa Rica; William M. Gabb collector]. Brocchi, Mission + scientifique au Mexique ..., pt. 3, sec. 2, Etudes 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. Guenther, Biologia Centrali-Americana: Reptilia + and Batrachia, p. 274, Sept. 1901. Werner, Abhand. Konigl. Akad. + Wissen. Muenchen., 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, Etudes 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. Guenther, 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]. Guenther, 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.) + +_Description and variation._--Ten breeding males from 15 to 20 +kilometers west-southwest of San Isidro el General, San Jose, Costa +Rica, have snout-vent lengths of 38.1 to 42.6 mm. (40.5 mm.). In these +specimens, the tibia/snout-vent length ratio is 0.50 to 0.54 (0.52), and +the tympanum/eye ratio is 0.45 to 0.57 (0.49). Specimens from the +Pacific slopes of Costa Rica are larger than those from the Meseta +Central and the Caribbean lowlands. Ten males from 6 kilometers east of +Golfito, Puntarenas, have snout-vent lengths of 38.4 to 44.6 mm. (41.8 +mm.), and five males from Rincon, Peninsula de Osa, have snout-vent +lengths of 38.8 to 41.6 mm. (40.3 mm.). Snout-vent lengths of ten males +from La Fortuna, Alajuela, are 31.9 to 36.0 mm. (34.4 mm.), of ten males +from Pandora, Limon, 33.8 to 37.6 mm. (35.9 mm.), and of ten males from +Escazu and Rio Jorco on the Meseta Central, 34.3 to 37.6 mm. (36.0 mm.). +Eight females from the Rio Jorco on the Meseta Central have snout-vent +lengths of 48.8 to 53.8 mm. (50.4 mm.), and six females from various +localities on the Pacific slopes of Costa Rica have snout-vent lengths +of 56.5 to 64.0 mm. (59.8 mm.). The only noticeable differences in +proportions between males and females is in the tympanum/eye ratio; for +example, this ratio is 0.47 to 0.53 (0.49) and 0.54 to 0.68 (0.61) in +ten males and eight females, respectively, from the Meseta Central. + +The shape of the snout and the associated cranial elements of _S. +sordida_ vary geographically and ontogenetically. Specimens from the +Caribbean lowlands have blunt snouts in lateral view; those from the +Pacific lowlands have longer, more slender snouts that are pointed in +lateral view, and those from the Meseta Central are intermediate in +snout shape between the two lowland populations (Fig. 4). These +differences in shape of the snout are dependent on the nature of the +underlying cranial bones, principally the maxillaries and nasals. In +specimens from the Caribbean lowlands the nasals are long, wide, and +narrowly separated from the ethmoid; the anterior edge is just posterior +to the nostril. The maxillary flanges are nearly vertical. In specimens +from the Pacific lowlands the nasals are relatively shorter, narrower, +and rather widely separated from the ethmoid; the anterior edges of the +nasals do not extend so far forward as in specimens from the Caribbean +lowlands. The maxillary flanges slant medially. In these cranial +characters, specimens from the Meseta Central are intermediate between +the two lowland populations. + +Superimposed on this geographic variation are ontogenetic changes, which +are most noticeable in males. In smaller, and presumably younger, +specimens the snouts are more pointed than in larger specimens; +consequently some small males from the Caribbean lowlands resemble +larger males from the Pacific lowlands, since the nasals and maxillaries +of the former are not fully ossified. In addition, in small breeding +males the ethmoid is only about one-half ossified, a large +frontoparietal foramen is present, the anterior arm of the squamosal +extends only about one-fourth the distance to the maxillary (two-thirds +the distance in larger specimens), and the tegmen tympani are short, as +compared with the long, thin elements in larger specimens. + + [Illustration: FIG. 4. Variation in the shape of the snout in + _Smilisca sordida_; left column females, right column males; + all from Costa Rica: (A) Camp Seattle, Rincon de Osa, Puntarenas + Prov. (UMMZ 123684); (B) Quebrada Agua Buena, 3 km. SW Rincon de + Osa, Puntarenas Prov. (USC 7236); (C) Rio Oro, 28.5 km. NW Villa + Neily, Puntarenas Prov. (KU 91742); (D) Rio Jorco, near + Desamparados, San Jose Prov. (KU 91765); (E-F) Bambu, Limon Prov. + (USC 7183). x3.] + +The dorsal ground-color of _Smilisca sordida_ is gray to pale tan or +reddish brown; the venter is white. The dorsum is variously marked with +dark gray, dark brown, reddish brown, or olive-green spots or blotches +(Pl. 7C). A dark interorbital bar usually is present. The dorsal +markings on the body usually consist of a blotch, or two or more spots, +on the occiput, in the scapular region, and in the sacral region. In +many specimens, especially females, these markings are in the form of +broad transverse bars. A female (USC 7164) from Las Canas, Guanacaste, +Costa Rica, has a tan dorsum with many black flecks and round brown +spots bordered by darker brown. One female (KU 91763) from the Rio +Jorco, San Jose, Costa Rica, has a unicolor tan dorsum. Some individuals +have scattered, small white spots on the dorsum; these are most evident +in a male (USC 7153) from La Fortuna, Alajuela. White labial stripes and +anal stripes are absent in all specimens. + +The limbs are marked by dark brown transverse bars; these are indistinct +in some specimens from the Meseta Central and Caribbean lowlands, +whereas they are distinct in all specimens from the Pacific lowlands. +Specimens from the Caribbean lowlands have two to six bars on each +shank, whereas specimens from the Pacific slopes have four to six bars +on each shank, and specimens from the Meseta Central have as many as +eight bars on each shank. A narrow, sometimes broken white line is +present on the ventrolateral edge of the forearm. The webbing on the +hand is tan or pale gray, and the ventral surfaces of the tarsi and the +webbing on the feet are dark gray or brown. Breeding males have dark +brown nuptial excrescences on the prepollex. + +The flanks and posterior surfaces of the thighs usually are marked by +bluish white and creamy tan flecks, respectively, but vary considerably. +In specimens from the Caribbean lowlands a small amount of flecking is +present in the inguinal region, and on the posterior surfaces of the +thighs flecks are few or absent. In specimens from the Meseta Central, +numerous large flecks or small, round spots (pale bluish white in life) +are on the posterior half of the flanks; small flecks are on the +posterior surfaces of the thighs. Specimens from the Pacific slopes and +lowlands of southern Costa Rica (Puntarenas and San Jose Provinces) have +bold mottling of black and bluish white on the flanks and many bluish +white flecks on the posterior surfaces of the thighs. The flanks are +reticulated from the axilla to the groin in two females (UMMZ 123684 and +USC 7236) from Rincon, Peninsula de Osa. In specimens from the Pacific +slopes of Guanacaste in northwestern Costa Rica, flecks are present in +the inguinal region; indistinct flecks are on the posterior surfaces of +the thighs. + +The throat is immaculate in specimens from the Caribbean lowlands in +Limon Province; the throats are dusky laterally in most other specimens +except some from the Meseta Central, in which the throats are heavily +flecked with black. This variation occurs in males and females. + +The color and pattern in life are highly variable. A composite +description of living individuals (now KU 91718-41) from 6 kilometers +east of Golfito, Puntarenas, Costa Rica, illustrates the variability: +"Dorsum pale olive-green, fading to tan posteriorly, or tan all over +with dark olive-green or dark brown spots on back and bars on limbs. +Flanks dark brown with cream, greenish gray, or bluish gray mottling. +Posterior surfaces of thighs dark brown with pale blue, pale green, or +tan flecks. Iris creamy silver. Throats white with some brown flecks +peripherally." (Duellman, Field notes, February 15, 1965.) A male from +the Rio Jorco, San Jose, Costa Rica, was dull olive-tan above with +olive-green marks; the flanks were brown with pale tan flecks, and the +posterior surfaces of the thighs were pale brown with cream-colored +flecks. Six females from the same locality were reddish brown above with +olive-brown or dark brown markings; one was uniform orange-tan, and +another was dull olive-green with darker markings. + +The color of the iris in living frogs varies from creamy silver to +grayish yellow or bronze with a variable amount of black reticulation. + +_Natural History._--_Smilisca sordida_ is not associated with any one +type of vegetation; instead it lives in the vicinity of rocky streams +having low gradients. Breeding takes place primarily in the dry season, +when the water in the streams is clear and at a low level. Through most +of the range of _S. sordida_ showers, or even short heavy rains, occur +in the dry season. After such rains the breeding activity is maximal. +Breeding congregations have been found from December through April, but +a few calling males and gravid females have been taken in June, July, +and August. In the rainy season non-breeding individuals are found +sitting on bushes near streams at night. Taylor (1952:843) found +specimens in bromeliads by day. + +Males usually call from rocks or gravel bars in, or at the edge of, +streams. Some individuals perch in low bushes overhanging the streams, +and some sit in shallows in the streams. Clasping pairs have been found +on the banks of streams and in shallow water in streams. + +The breeding call consists of one to six moderately short, rather +high-pitched notes (duration 0.18 to 0.45 seconds) repeated at intervals +of 12 seconds to several minutes. Each note is a vibrant rattle having +78 to 135 pulses per second and major frequences of about 1200 to 2600 +cycles per second (Pl. 11C). + +The tadpoles live in shallow parts of the streams, where they cling to +the surfaces of small rocks and hide beneath leaves and rocks. A +complete developmental series of tadpoles is not available; measurements +of those stages examined are summarized in Table 12. + +A typical tadpole in stage 36 of development (KU 68475 from 15 km. WSW +of San Isidro el General, Costa Rica) has a body length of 11.7 mm., +tail length of 22.8 mm., and a total length of 34.5 mm.; body about +three-fourths as deep as wide; snout broadly rounded in dorsal view, +sloping and rounded in lateral view; eyes widely separated, directed +dorsolaterally; nostril slightly closer to eye than to tip of snout; +mouth ventral; spiracle sinistral, about two-thirds distance from snout +to posterior end of body and slightly below midline; anal tube dextral; +caudal musculature heavy, straight; dorsal fin not extending onto body; +fins deepest at about mid-length of tail; there depth of caudal +musculature equal to depth of dorsal fin and half again as deep as +ventral fin; musculature extending nearly to tip of tail; body reddish +brown above and pale grayish brown with white flecks below; caudal +musculature pale tan with brown flecks; a series of reddish brown dashes +at base of caudal fin separated from others in series and from dashes on +other side by creamy white; fins transparent with reddish brown flecks +on posterior one-half of ventral fin and on all of dorsal fin (Fig. +14C). Mouth bordered by two rows of short, pointed papillae; lateral +fold present; tooth-rows 2/3; upper rows equal in length; second upper +row narrowly interrupted medially; three lower rows complete, nearly as +long as upper rows, deeply indented medially; upper beak robust, inner +surface not forming continuous arch with short lateral processes; lower +beak deep, V-shaped; both beaks bearing short serrations (Fig. 15F). + +Little variation occurs in structure. In some specimens the second upper +tooth-row is complete; no individuals were found to have the row broadly +interrupted medially. + +The series of dark dashes on the dorsal edge of the caudal musculature +is diagnostic of all stages studied. In life, tadpoles from 15 and 20 +kilometers west-southwest of San Isidro el General, Costa Rica, had a +tan body, often with an olive-tan tinge; the caudal musculature was +tan; the flecks and dashes were dull red or reddish brown. Tadpoles from +6 kilometers east of Golfito, Costa Rica, had bodies with olive-green +flecks. The caudal musculature was brown with bluish green flecks; the +fins were transparent with reddish brown flecks. The belly was a silvery +golden color. Tadpoles from Bajos de Jorco, Costa Rica, had brown bodies +with bluish green flecks; the tail and fins had reddish brown flecks and +dashes. The iris was a bronze color in specimens from all three +localities, as well as in the young mentioned in the following +paragraph. + +Nine recently metamorphosed young were found on vegetation at the edges +of streams in April. These specimens have snout-vent lengths of 13.1 to +15.7 mm. (14.9 mm.) and in life were pale greenish tan or olive-tan +above and white below. The hands, feet, and thighs were pale yellowish +tan. + +_Remarks._--The foregoing synonymies indicate that confusion has existed +in the application of various names, to this species, as well as in use +of the names _sordida_ and _gabbi_ to include the species that we +describe and name _Smilisca sila_. Correct allocation of the names +involved was possible only after studying and comparing the type +specimens, for the descriptions given by the various authors are not +sufficiently explicit to determine the nature of many essential +features. + +The presence of a rounded snout and a long white throat in males +distinguishes _S. sordida_ from _S. sila_, which has a high truncate +snout and short dark throat in males. The two syntypes of _Hyla sordida_ +Peters, 1863, (ZMB 3141) are males having snout-vent lengths of 36.9 and +37.0 mm. The two syntypes of _Hyla gabbi_ Cope, 1876 (USNM 30658-9), are +females having snout-vent lengths of 52.8 and 53.7 mm., respectively. +Also included in the collections made by Gabb is eastern Costa Rica are +two males (USNM 30685-6), which Cope (1876) named and described as _Hyla +nigripes_. These specimens are soft and faded, but are recognizable as +the same as _Hyla sordida_ Peters; the syntypes of _Hyla nigripes_ have +snout-vent lengths of 37.6 and 37.7 mm. We have examined one of the +syntypes of _Hyla salvini_ Boulenger, 1882 (BMNH 1947.2.24.13), a female +having a snout-vent length of 54.6 mm. We are convinced that all of +these type specimens are representatives of one species, the earliest +name for which is _Hyla sordida_ Peters, 1863. The type localities for +three of the named species are in Costa Rica--_H. gabbi_ from Sipurio on +the Caribbean lowlands, _H. nigripes_ from the Caribbean slopes of Pico +Blanco, and _H. salvini_ from Cartago on the Meseta Central. The type +locality of _H. sordida_ was given as "Veraguas" by Peters (1863). At +that time Veraguas was often considered to be most of western Panama. +Though we have not seen Panamanian specimens other than the types of _S. +sordida_ and one specimen from the Pacific lowlands of western Panama, +the species probably occurs on the Caribbean slopes of western Panama. +The species has been taken on the Caribbean lowlands of Costa Rica +within a few kilometers of Panama; collecting on the Caribbean slopes in +the provinces of Bocas del Toro and Veraguas should reveal the presence +of _Smilisca sordida_ there. + +_Distribution._--_Smilisca sordida_ is found along the Pacific slopes +and lowlands from Guanacaste, Costa Rica, southeastward to extreme +western Panama, to elevations of about 1200 meters on the Meseta Central +in Costa Rica, and on the Caribbean slopes and lowlands of Costa Rica +and probably adjacent Panama (Fig. 5). One specimen purportedly comes +from "Rio Grande, Nicaragua." + + [Illustration: FIG. 5. Map showing locality records for + _Smilisca sordida_.] + +_Specimens examined._--412, as follows: NICARAGUA: "Rio Grande" +(? Depto. Zelaya), MCZ 2634. + +COSTA RICA: =Alajuela=: Between Atena and Salto de San Mateo, USC 6185; +8 km. N Ciudad Quesada, USC 7155 (4); La Fortuna, USC 7153 (20); 3 km. E +La Fortuna, USC 7150; San Carlos, USNM 29969; Sarchi, KU 32990-9, +36792-3. + +=Cartago=: Cartago, BMNH 1947.2.24.13; headwaters of Rio Pacuare, USC +119; Instituto Interamericano de Ciencias Agricolas, Turrialba, KU +37012, USC 420, 437; Rio Reventazon, Turrialba, MCZ 29268: 10 km. N Rio +Reventazon bridge, USC 7073; 5 km. SW Rio Reventazon bridge on +Paraiso-Orosi road, USC 669; Turrialba, UMMZ 118405, USC 455, USNM +29936-9. + +=Heredia=: Puerto Viejo, KU 36791. + +=Guanacaste=: Las Canas, USC 7164; Santa Cecilia, MCZ 7924-5; Tilaran, +USC 7161 (5). + +=Limon=: Bambu, USC 7171 (2), 7183 (13); La Lola, USC 820 (6), 6083-94, +8064, 8071; Pandora, USC 7188 (7), 7189, 7190 (3), 7191 (5); Pico +Blanco, USNM 30685-6; Rio Lari, 14-16 km. SW Amubre, USC 7179, 7180 +(10); Sipurio, USNM 30658-9; Suretka, KU 36764, 36765 (skeleton), +36766-78. + +=Puntarenas=: 6 km. N Dominical, KU 91749-50, 91811 (young), 91812 +(tadpoles); Esparta, MCZ 8028; 6 km. E Golfito, KU 91718-41, 91809 +(young), 91810 (tadpoles), 91816-9 (skeletons), USC 7103 (23); Quebrada +Agua Buena, 3 km. SW Rincon de Osa, USC 7236 (6); Quebrada Boruca, 22 +km. E Palmar Norte, KU 64264; Rincon de Osa, Camp Seattle, UMMZ +123680-5, S-2792 (skeleton), USC 705 (5), 6023, 7254; Rio Barranca, USC +7119 (2); Rio Ceiba, 6 km. NW Buenos Aires, KU 91747-8, USC 7112 (7); +Rio Ciruelitas, 16 km. NW Esparta, USC 7121 (3); Rio Claro, 14.2 km. NW +Villa Neily, USC 7110 (4); Rio Ferruviosa, 7 km. S Rincon de Osa, USC +7235 (4); Rio Lagarto at Pan-American Hwy. (Guanacaste Border), USC 7122 +(4); Rio La Vieja, 30 km. E Palmar Norte, KU 87684 (tadpoles), 91743-6, +USC 7083 (2); Rio Oro, 28.5 km. NW Villa Neily, KU 91742; Rio Volcan, 10 +km. W Buenos Aires, USC 7113; Rio Zapote, 7 km. E Palmar, USC 7100 (4); +3-5 km. W Palmar, USC 7101 (18); 7 km. SE Palmar, KU 64261-3; 1.2 km. NW +Villa Neily, USC 8032; 3 km. NW Villa Neily, USC 7109 (20); 5 km. NW +Villa Neily, USC 6176, 8035. + +=San Jose=: Bajos de Jorco, KU 91813 (tadpoles); Escazu, KU 34863, +34869-75, USC 813; between Monrovia and La Hondura, +- 0.5 km. N Santa +Rosa, USC 302 (2); Paso Ancho, Rio Jorco, UMMZ 122649 (6), USC 530 (3); +Rio Jorco, near Desamparados, KU 91757-65, 91796-7, 91820-3 (skeletons), +USC 228, 513, 7117 (7); Rio Peje, 10 km. SSE San Isidro el General, USC +7115 (3); Rio Tirivi, MCZ 7972; San Isidro el General, CNHM 101096, KU +28201, 32989, UMMZ 72024; 15 km. WSW San Isidro el General, KU 64245-56, +68473 (tadpoles), 68474 (young), 68475 (tadpoles), 86516, 91754-6, +91793-5, USC 7097 (6); 17.1 km. WSW San Isidro el General, USC 6047; 18 +km. WSW San Isidro el General, USC 689; 20 km. WSW San Isidro el +General, KU 64257-9, 64260 (skeleton), 68468 (young), 68469 (tadpoles), +68470 (young), 68471-2 (tadpoles), 68476 (young), 68633-4 (skeletons), +91751-3; San Jose, AMNH 7501-4, USC 298; Santa Rosa, Rio Virilla, USC +7145. + +PANAMA: =Chiriqui=: Rio Jacu, 5.8 km. ESE Paso Canoas, KU 91905. +"Veraguas," ZMB 3141 (2). + + + + +ANALYSIS OF MORPHOLOGICAL CHARACTERS + + +Osteology + +In attempting to assay the taxonomic significance of skeletal +differences we are faced with a dearth of data on the skeletons of frogs +in general and hylids in particular. Recent reviews by Brattstrom (1957) +and Hecht (1962, 1963) have been concerned with general salientian +classification and phylogeny, principally at the family level. Savage +and Carvalho (1953), Griffiths (1959), and Baldauf (1959) used +osteological characters in determining the taxonomic status of the +families Pseudidae, Brachycephalidae, and Bufonidae, respectively. +Carvalho (1954) presented osteological evidence for the generic +separation of New World microhylids. Zweifel (1956) and Tihen (1962) +used osteological characters at the levels of the species-group and +species in their respective studies on _Scaphiopus_ and _Bufo_. Little +has been recorded about the skeletons of the hylids. Goin (1961) +mentioned dentigerous elements and cranial co-ossification in his +synopsis of the genera of hylids. Copland (1957) in his review of +the _Hyla_ of Australia, Funkhouser (1957) in her revision of +_Phyllomedusa_, and Zweifel (1958) in his review of _Nyctimystes_ did +not consider skeletal characters. + +Some osteological studies on hylids have yielded worthwhile information. +Mittleman and List (1953) used osteological characters in defining the +genus _Limnaoedus_: Starrett (1960) used cranial characters in +combination with jaw musculature in defining the genus _Smilisca_, and +Duellman (1964) used cranial characters in delimiting the _Hyla +bistincta_ group. Brief descriptions of cranial structure were given for +_Phrynohyas_ (Duellman, 1956) and _Ptychohyla_ (Duellman, 1963a); +specific and sexual differences in the skulls of _Hyla chaneque_ and +_Hyla taeniopus_ were pointed out by Duellman (1965). Stokely and List +(1954) described early cranial development in the hylid _Pseudacris +triseriata triseriata_. + +Because our knowledge of the skeleton in hylids is so incomplete, we are +not attempting to place _Smilisca_ in the general scheme of hylid +phylogeny on the basis of skeletal characters. Instead, our purposes are +to describe the skeleton and its ontogenetic development in one member +of the genus (_S. baudini_), and to make comparisons that show taxonomic +differences in osteological characters among species of _Smilisca_. + +The study of 68 dried skeletons and 25 cleared and stained preparations, +including an ontogenetic series of _S. baudini_, has resulted in an +understanding of the progressive development of skeletal elements and a +knowledge of interspecific and intraspecific variation in these +elements. Furthermore, investigations of the osteology have provided +correlations between some cranial characters and certain aspects of +external morphology. + +_Descriptive Osteology of Smilisca baudini_ + +The following description is based primarily on an adult female (KU +68184): + +_Skull._--The skull is large, solid, and broader than long; the greatest +width is between the sutures of quadratojugal and maxillary on either +side of the skull (Pls. 2-3). The maxillaries bear well-developed dorsal +flanges, curve gently, join the moderately convex premaxillaries +anteriorly and form a slightly truncate snout. The combined premaxillary +width is about one-fourth the width of the skull. The premaxillaries are +separated medially, and laterally from the maxillaries by sutures. Each +premaxillary bears a dorsomedial alary process, which is anteriorly +convex and four times as high as the depth of the lateral wing of +premaxillary; each premaxillary also has a ventromedial palatine process +that projects dorsally from the lingual edge of the premaxillary. The +septomaxillaries are closely associated dorsally with the premaxillaries +immediately lateral to the prenasal processes. + +The nasals are large, widest anteriorly and narrowing posteriorly, +parallel to maxillaries, and not separated from the ethmoid by +cartilage. The nasals bear long, delicate maxillary processes extending +nearly to the maxillaries. Anteriorly, the nasals are widely separated +by the partially ossified internasal septum, which is in contact with +the premaxillaries between the prenasal processes; the anterior points +of the nasals lie approximately one-half the distance between the +anterior ends of the ethmoid and the premaxillaries. The ethmoid is +large and completely ossified; the margins are smooth. The trunate +anterior edge lies between the nasals and is in contact with the +internasal septum. The frontoparietals are large, smooth-margined, and +bear large supraorbital flanges curving posterolaterally at the rear of +the orbit. A small, oval foramen involves the posterior part of the +ethmoid and anterior portion of frontoparietals; continued ossification +in older specimens fills in the foramen, thereby resulting in a solidly +roofed cranium. The auditory regions are relatively massive and bear +narrow tegmen tympani; the distal ends of the tegmen tympani are medial +to the lateral edge of the pterygoids in dorsal view. The squamosals are +large; the long anterior arm is separated from the maxillary by a +suture. The delicate, spindle-shaped columellae lie ventral to the +tegmen tympani and squamosals, are spatulate distally, and have a broad +basal attachment to the auditory region. + +The vomers are moderately large and are in contact anteriorly with the +premaxillaries and posteriorly with the ethmoid. Each vomer has two wide +serrated flanges laterally. The tooth-bearing parts of the vomers are +widely separated and at a slight angle to one another; the vomers +terminate medially in two pointed processes on the ethmoid. The +palatines are edentate, but bear strong ridges throughout their lengths. +They are broadly in contact with the maxillary, are narrow medially, and +are attached by pointed processes to the medial part of the ethmoid. +The pterygoids are large, attached to the maxillaries immediately +anterior and medial to the squamosal-maxillary connection, bear +well-developed pedicles, which are broadly attached to the prooetic, and +a wide wing is in contact posteriorly with the distal two-thirds of the +quadrate. + +The angular makes up most of the lower jaw, bears a broad articular +surface posteriorly, and has a small coronoid process on the lingual +edge; anteriorly the angular is separated from the dentary and +mentomecklian by Meckel's cartilage. The dentary lies external to the +angular and extends from the mentomecklian to approximately the +mid-length of the angular. The mentomecklians are ossified, but +separated by cartilage medially. + +_Hyoid._--The hyoid plate is curved, thin, and mostly cartilaginous, but +calcined posteriorly (Fig. 6). The anterior cornua are slender, +cartilaginous, and curve anteromedially from the hyoid plate and thence +laterally and posteriorly, to attach to the posterior surface of the +prooetics. The lateral cornua are broad, flat, cartilaginous lateral +extensions from the bases of the anterior cornua. The posterior cornua +are bony, except distally. + + [Illustration: FIG. 6. Ventral view of hyoid apparatus of an adult + male _Smilisca baudini_ showing areas of muscle attachment: _Gen. + L._, attachment of geniohyoideus lateralis; _Gen. M._, attachment + of geniohyoideus medialis; _Hyo._, attachment of hyoglossus; _Omo._, + attachment of omohyoideus; _Pet._, petrohyoideus; _St._, attachment + of sternohyoideus. KU 64220, x5.] + +_Vertebral Column._--The atlas lacks transverse processes and a neural +crest, whereas transverse processes are present on the other seven +presacral vertebrae, and knoblike neural crests are present on the +second, third, and fourth vertebrae; a faint neural ridge is visible on +the fifth vertebra. The transverse processes are directed laterally on +the second and sixth vertebrae, ventrolaterally on the third, +posterolaterally on the fourth and fifth, and anterolaterally on the +seventh and eighth. The processes are slightly expanded on the fourth, +and more so on the fifth, vertebra. The sacral diapophyses are expanded +and have a border of calcified cartilage laterally. There are two sacral +condyles. The slender coccyx has a thin dorsal ridge on the anterior +three-fourths of its length. + +_Pectoral Girdle._--The omosternum is large, ovoid, and cartilaginous; +the sternum is a thin cartilaginous sheet deeply notched posteriorly and +is not differentiated into episternal and xiphisternal elements. The +coracoids are robust, twice as stout as the clavicles. The epicoracoidal +cartilages overlap in the usual arciferal manner, except that they are +fused anteriorly between the slender clavicles. The clavicles are +strongly arched. The clavicle, coracoid, and scapula on each side form a +bony articulation at the glenoid fossa. A bifurcation of the ventral end +of the scapula results in a large glenoid foramen. The scapula is flat +and expanded dorsally; the suprascapula is broad, flat, and calcified in +large adults. In young specimens no distinct ossification of the +cleithrum or ossification of endochondral centers are evident. + +_Arm and Hand._--The humerus is equally well-developed in both sexes and +has a prominent lateral crest. The radius and ulna are completely fused. +A bony prepollex is present in both sexes. The metacarpals are about +equal in length. The phalangeal formula is 2-2-3-3; the terminal +phalanges are claw-shaped. + +_Pelvic Girdle._--The ilia are long, slender, and slightly curved. A +thin ridge projects laterally from the dorsal edge of the posterior +one-half of each ilium. The ilial prominence is large and knoblike when +viewed from above. The anterior edge of the ilial prominence is at the +level of the anterior edge of the acetabular border. The dorsal +acetabular expansion is small. The pubis is slender, and the ischium is +elevated and robust. + +_Leg and Foot._--The slightly curved femur has a distinct crest +proximally on the posterior surface. The nearly straight tibio-fibula is +slightly longer than the femur. The tibial and fibial elements are +completely fused but have a distinct cleft between them. A small foramen +exists at the mid-length of the tibio-fibula. The fibulare (calcaneum) +is much more robust than the tibiale (astragalus). The prehallux is +large and flat. The metatarsals of the third, fourth, and fifth digits +are equal in length; the metatarsal of the second is somewhat shorter, +and that of the first is much shorter. The phalangeal formula is +2-2-3-4-3; the terminal phalanges are claw-shaped. + +_Developmental Cranial Morphology of Smilisca baudini_ + +The following description of development of the skull of _Smilisca +baudini_ is based on the examination of 12 cleared and stained +specimens. In table 3 the cranial bones are listed in the left hand +column in the approximate order of their appearance in the young frogs. +Across the top of the table selected specimens designated by +developmental stage or snout-vent length are listed. It should be noted +that although each individual, from left to right, has an increasing +number of ossified bones, the correlation with increasing size is +imperfect; the precise ages of the individuals are unknown. + +The first bones to appear are the septomaxillaries, frontoparietals, +part of the exoccipital, and the parasphenoid in developmental stage 40. +The frontoparietals are represented by two slender ossifications +dorsomedial to the orbits; the septomaxillaries are present as small +ossifications anterior to the nasal capsules (Pl. 1A). The parasphenoid +is present as a faint median ossification, and the exoccipital shows +some ossification. + + + Table 3.--The Order of Occurrence of Cranial Ossifications in the + Skull of Smilisca baudini. Where Numbers Are Divided by a Slash + Mark, the Left and Right Symbols Correspond to the Left and + Right Sides of the Skull, Respectively. + + =====================+=====+=====+=====+=====+=====+=====+==== + Bone |Stage|Stage|12.6 |13.9 |32.0 |27.0 |20.1 + | 40 | 44 | mm. | mm. | mm. | mm. | mm. + ---------------------+-----+-----+-----+-----+-----+-----+---- + Frontoparietal | X | X | X | X | X | X | X + Parasphenoid | X | X | X | X | X | X | X + Septomaxillaries | X | X | X | X | X | X | X + Exoccipitals | X | X | X | X | X | X | X + Squamosals | -- | X | X | X | X | X | X + Premaxillaries | -- | X | X | X | X | X | X + Maxillaries | -- | X | X | X | X | X | X + Nasals | -- | -- | X | X | X | X | X + Pterygoids | -- | -- | X | X | X | X | X + Vomers | -- | -- | -- | X | X | X | X + Palatines | -- | -- | -- | X | X | X | X + Quadratojugals | -- | -- | -- | X | X | X | X + Ethmoid | -- | -- | -- | -- | X | X | X + Columellas | -- | -- | -- | -- | X | X | X + Supraorbital Flanges | -- | -- | -- | -- | -- | X | X + Prooetics | -- | -- | -- | -- | -- | -- | X + Vomerine Teeth | -- | -- | 1/1 | 4/3 | 5/5 | 3/3 | 5/4 + Maxillary Teeth | -- | 0/7 | 3/5 | 6/5 |30/31|30/26|37/36 + Premaxillary Teeth | -- | 2/4 | 3/3 | 5/5 | 7/6 | 8/6 | 8/7 + ---------------------+-----+-----+-----+-----+-----+-----+---- + + +The dentigerous bones are among the most rapidly developed, although not +the first to appear. They are present in developmental stage 44 before +metamorphosis is completed. The maxillaries bear a few teeth anteriorly +and are ossified posteriorly to a point one-third of the distance from +the anterior to the posterior edge of the orbit. Ossification lengthens +the posterior termini of the maxillaries to the posterior edge of the +orbit. In front of the anterior margin of the orbit, bone is +proliferated dorsal to the main axes of the maxillaries and forms +moderate dorsal maxillary flanges. The premaxillaries appear +simultaneously with the maxillaries. Initially they are widely separated +medially from each other, and laterally from the developing maxillaries; +each bears two or three teeth, large dorsally blunt alary processes, and +small palatine processes. The median and lateral edges of the prenasal +processes lengthen heterochronously, causing the median edges to be +longest and to lie slightly dorsal to the level of the septomaxillaries. +After the maxillaries and premaxillaries develop, the vomers appear as +small horizontal ossifications anterior to the parasphenoid. +Ossification begins in the lateral flanges, then in the prevomerine +processes, and lastly in the posterior dentigerous parts of the bones; +the prevomerine processes are the last parts of the vomers to ossify +completely. + +Initially the frontoparietals are present as thin rods of ossification +dorsomedial to the orbits; the frontoparietals extend from the anterior +to the posterior end of the orbit by developmental stage 44. The +anterior ends of the bones remain thin and pointed; ossification +progresses medially from the midpoint of the length of the orbit and +posteriorly to the level of the exoccipital; a median center of +ossification joins the frontoparietals posteriorly, thereby forming the +posterior border of the frontoparietal fontanelle. The supraorbital +flanges of the frontoparietals do not appear until all other cranial +bones are ossified, or nearly so. The most rapid ossification begins +laterally at the posterior edge of the orbit and decreases anteriorly +over the posterior half of the orbit. This differential rate of +proliferation of bone results in the pattern of development of the +supraorbital flanges shown in figure 7. The nasals appear as +thin slivers of bone half way between the anterior ends of the +frontoparietals and the end of the snout. As ossification proceeds the +nasals assume a triangular shape in dorsal view. The anterior ends are +pointed; the lateral margins are parallel to the maxillaries. The +posteromedial points do not reach the lateral margins of the ethmoid, +and the maxillary processes extend about three-fourths the distance from +the bodies of the nasals to the maxillaries. Following the union of the +frontoparietals posteriorly, the nasals widen anteriorly and are +narrower at the midpoints of their long axes than anteriorly or +posteriorly. With further ossification the maxillary processes extend to +the maxillaries and form complete bony anterior margins to the orbits; +the mid-parts of the nasals widen (Pl. 1B). + + [Illustration: FIG. 7. Developmental sequence of the frontoparietal + fontanelle and associated bony elements in _Smilisca baudird_: + (A) KU 60026, x5; (B) KU 85438, x4; (C) KU 26328, x3; + (D) KU 68184, x2.3.] + +The parasphenoid is the first of the palatal bones to appear. At +metamorphosis the bone is well developed; the anterior tip is situated +just in front of the anterior edge of the orbit, and posteriorly the +lateral processes extend laterally beyond the ossified parts of the +auditory region. The pterygoids do not appear until metamorphosis, when +ossification is evident in only the mid-parts of the posterolateral +arms. Ossification follows in the mid-parts of the anterolateral arms +and occurs last in the pterygoid pedicles. The palatines do not appear +until all three arms of the pterygoids are at least partly ossified. +Ossification proceeds rapidly from the maxillaries medially to the +unossified ethmoid, which is the last of the cranial bones to appear. +Initially it is extremely shallow; dorsally it is widely separated from +the nasals, and ventrally the posterior margin meets the anterior point +of the parasphenoid. In dorsal view, ossification proceeds anteriorly +between the nasals and posteriorly, ventral to the frontoparietals; +ventrally, ossification proceeds posteriorly dorsal to the parasphenoid. + +The ventral arms of the squamosal and the supraoccipital region of the +exoccipital are the first occipital bones to appear. Ossification +follows in the regions of the semicircular canals and occipital +condyles. The dorsal end of the ventral arm of the squamosal and the +posterior arm of the squamosal ossify as a unit at the same time the +quadratojugal appears. Shortly thereafter the anterior arm of the +squamosal ossifies, the distal part of the columella appears, and the +anterior and lateral parts of the auditory region ossify. + +The angular and dentary of the lower jaw appear concurrently with the +dentigerous bones. Initially, the angular is short and broad; the +articular surface is absent, and the anterior end is slightly overlapped +by the dentary. The mentomecklians do not ossify until approximately +the same time that the quadratojugal appears in the upper jaw. + +_Comparative Osteology_ + +The genus _Smilisca_ is characterized by the following combination of +cranial osteological characters: (1) A large amount of bone is involved +in the skull and a minimal amount of cartilage and/or secondarily +ossified cartilage; co-ossification is absent. (2) The skulls are +uniformly broad with angular lateral margins, and truncate anteriorly. +(3) An internasal septum and quadratojugals are present. (4) A +well-developed squamosal minimally extends one-fourth the distance from +the dorsal end of the quadrate to the maxillary, and maximally is +separated from the maxillary by a suture. (5) The ethmoid is large; the +distance between the anterior end of the ethmoid and the anterior edge +of the premaxillary varies between 15 and 20 per cent of the total +length of the skull. + +On the basis of cranial osteology two species-groups can be recognized +within the genus _Smilisca_. The _sordida_ group, comprising _S. +sordida_ and _puma_, is characterized by a broad skull in which the +lateral margins of the maxillaries are relatively straight anterior to +the orbit. The moderate-sized nasals are rounded anteriorly, and bear +relatively short, sometimes blunt, maxillary processes. The long axes of +the nasals are not parallel to the maxillaries. The ethmoid is +proportionately small in the _sordida_ group. The bony part of the +ethmoid terminates near the anterior edge of the orbits and does not +extend anteriorly between the nasals; the entire anterior margin of the +ethmoid is separated from the nasals by cartilage. The squamosals are +generally small. They are narrow in dorsal view, and minimally extend +one-fourth the distance from the dorsal end of the quadrate to the +maxillary, and maximally, two-thirds the distance. The tegmen tympani +are relatively small (Fig. 8). + + [Illustration: FIG. 8. Dorsal views of the skulls of the species of + _Smilisca_: (A) _S. baudini_ (KU 68184); (B) _S. puma_ (KU 68636); + (C) _S. phaeota_ (KU 41090); (D) _S. sila_ (KU 80625); (E) _S. + cyanosticta_ (KU 55938), and (F) _S. sordida_ (KU 36765). x1.5.] + +In contrast to the tendency for reduction of cranial parts in the +_sordida_ group, the _baudini_ group, constituted by _S. cyanosticta_, +_phaeota_, and _baudini_, is characterized by more ossification of the +cranial elements. The skull is broad; the lateral margins are less +angular and are gently curved, rather than straight as in the _sordida_ +group. The nasals tend to be larger with the long axes parallel to the +maxillary. Anteriorly the nasals are pointed, and posteriorly they bear +long, delicate palatine processes extending to the maxillary. The +ethmoid is fully ossified, extends anteriorly between the nasals, and +laterally is separated by a suture from the nasals if the latter are +fully ossified. The squamosals are large, and wide in dorsal view. They +minimally extend one-fourth the distance from the dorsal end of the +quadrate to the maxillary, and maximally are sutured to the maxillary. +The tegmen tympani are massive. + +_Smilisca sila_ is intermediate between the two species-groups +described. The skull is broad; the lateral margins are gently curved, +and have a pronounced angularity just anterior to the palatines which +results in a broad, truncate snout. The nasals are moderate in size; +because of the anterior angularity of the lateral margins, the long axes +of the nasals lie parallel to the maxillary. The nasals are only +slightly pointed anteriorly, and posteriorly they bear short, blunt +palatine processes and medial processes in contact with the lateral +corners of the ethmoid. The ethmoid is fully ossified, but does not +extend anteriorly between the nasals. The squamosals are moderate in +size and extend one-fourth the distance from the dorsal end of the +quadrate to the maxillary. The tegmen tympani are relatively large, but +proportionately short. + +The cranial characters utilized in the analysis of species groups +(general shape, nature of the nasals, ethmoid, squamosals, and tegmen +tympani), together with other characters, such as the relative height +and shape of the prenasal processes, the extent of the internasal +septum, and the nature of the vomers, frontoparietals, maxillaries and +pterygoids are useful in distinguishing the various species (Table 4, +Fig. 8), as well as in establishing relationships within the +species-groups. + +Within the _sordida_ group, _S. sordida_ and _S. puma_ can be +distinguished by the following characters: The bony part of the ethmoid +terminates posterior to the anterior edge of the orbit and is thus +widely separated from the nasals by cartilage in _S. puma_. In _S. +sordida_ the bony part of the ethmoid always terminates at a level equal +to, or slightly in front of the anterior edge of the orbit; therefore, +less cartilage exists between the ethmoid and nasals in _S. sordida_ +than in _S. puma_. The width of the premaxillary comprises about 30 per +cent of the width of the skull in _S. sordida_ and 20 per cent in _S. +puma_. The proportion of the length of the skull anterior to the bony +part of the ethmoid in _S. sordida_ is approximately 21 per cent, as +compared with about 29 per cent in _S. puma_. The prenasal processes are +convex in _S. sordida_ and straight in _S. puma_. + +The marked ontogenetic variation in _S. sordida_ is considered in more +detail in the account of that species, but it is pertinent to the +present discussion to note that with respect to some features of the +skull some young breeding specimens of _S. sordida_ are intermediate in +appearance between large females of _S. sordida_ and adults of _S. +puma_. In some breeding males (usually the smaller individuals) of _S. +sordida_ the bony part of the ethmoid terminates at the anterior edge of +the orbit and is widely separated from the nasals by cartilage. In small +individuals _S. sordida_, especially in males, and in adults of _S. +puma_ the tegmen tympani are relatively short, whereas in adult females +of _S. sordida_ these elements are long and slender. In the smaller +specimens of _S. sordida_ and in _S. puma_ the squamosal is small; it +extends only about one-fourth of the distance to the maxillary in the +smaller _S. sordida_ and about one-half the distance in _S. puma_. The +more massive squamosal in large adult females of _S. sordida_ extends at +least two-thirds of the distance to the maxillary. + + + Table 4.--Comparative Cranial Osteology of Smilisca. + + ===============+==============================+======================= + Character | _S. baudini_ | _S. cyanosticta_ + ---------------+------------------------------+----------------------- + | | + Alary Processes| Four times as high as | Three times as high + | lateral wing of premaxillary;| as lateral wing of + | anteriorly | premaxillary; + | convex. | anteriorly + | | convex. + | | + Nasals | Long, wide anteriorly, | Long, widest + | narrowing posteriorly; | posteriorly; + | attached to ethmoid. | attached to + | | ethmoid. + | | + | | + | | + Ethmoid | Long; entirely ossified; | Long, entirely + | smooth margins. | ossified; + | | smooth margins. + | | + Frontoparietal | Small, ovid fontanelle | Large fontanelle, two + | present or absent; | and one-half times as + | long, pointed postorbital | long as wide; narrow + | processes curving | supraorbital flanges + | along posterior | with irregular margins. + | border of orbit. | + | | + Squamosal | Large: anterior arm | Large; anterior arm + | in contact with maxillary. | in contact with + | | maxillary. + ---------------+------------------------------+------------------------ + + TABLE 4 (Continued) + ===============+=============================+========================= + Character | _S. phaeota_ | _S. puma_ + ---------------+-----------------------------+------------------------- + | | + Alary Processes| Two and one-half | Two times as high as + | times as high as lateral | lateral wing of + | wing of premaxillary; | premaxillary; + | anteriorly convex. | straight. + | | + | | + Nasals | Long, widest anteriorly | Short, narrow, not + | and posteriorly, | attached to ethmoid. + | bearing posteromedial | + | process; not attached | + | to ethmoid. | + | | + | | + Ethmoid | Long, entirely ossified; | Short, about two-thirds + | smooth margins. | ossified; irregular + | | margins. + | | + Frontoparietal | Fontanelle absent; | Keyhole-shaped fontanelle; + | large supraorbital | smooth margins; + | flanges having | flanges absent. + | straight edges and extending| + | posterolaterally. | + | | + | | + Squamosal | Large; anterior arm | Small; anterior arm + | extending 1/2-2/3 way | extending 1/2 way to + | to maxillary. | maxillary. + ---------------+-----------------------------+-------------------------- + + TABLE 4 (Continued) + ===============+===========================+============================ + Character | _S. sila_ | _S. sordida_ + ---------------+---------------------------+---------------------------- + | | + Alary Processes| One and one-half | Two and one-half + | times as high as lateral | times as high as lateral + | wing of premaxillary; | wing of premaxillary; + | straight. | slightly convex + | | anteriorly. + | | + Nasals | Short, wide, bearing | Moderately long narrowest + | small posteromedial | anteriorly and + | processes; not attached | posteriorly; not attached + | to ethmoid. | to ethmoid. + | | + | | + | | + Ethmoid | Moderately long; entirely | Short; one-half to entirely + | ossified; smooth | ossified; irregular + | margins. | margins. + | | + Frontoparietal | Large, ovoid fontanelle; | Large, elongate fontanelle; + | smooth margins; | smooth margins; + | flanges absent. | flanges absent. + | | + | | + | | + | | + Squamosal | Moderately large; anterior| Moderately small; anterior + | arm extending | arm extending + | 1/4 way to maxillary. | 1/4-2/3 way to maxillary. + ---------------+---------------------------+---------------------------- + + +Within the _baudini_ group, the skull of _S. cyanosticta_ is the most +generalized of the three species; the cranial characters are +intermediate between _S. phaeota_ and _S. baudini_. The lateral margins +of the skull in _S. cyanosticta_ are gently curved, and have an +angularity anterior to the palatine-maxillary suture; the anterior +margins are less angular in _S. phaeota_, which has a broader snout. +Posteriorly in _S. baudini_ the margins are slightly curved +medially, and the greatest width of the skull is between the +quadratojugal-maxillary sutures on either side of the skull. The +frontoparietals of _S. cyanosticta_ bear slightly irregular lateral +margins and a large fontanelle. There is a tendency for obliteration of +the fontanelle with increasing age in both _S. baudini_ and _S. +cyanosticta_; the lateral margins of the frontoparietals bear large +supraorbital flanges in both of these species. In _S. phaeota_ the +flanges are most prominent; they extend posterolaterally with straight +margins along two-thirds of the length of the orbit and terminate in +rather blunt points. The broad interorbital flanges result in a +relatively broad external interorbital distance. In _S. baudini_ the +flanges are curved posterolaterally around the orbit and terminate in +sharp, thin points. The tegmen tympani of all three species are massive. +In _S. cyanosticta_ the prooetics slope posteriorly, whereas they slope +anteriorly in _S. baudini_ and _S. phaeota_. + +The skulls of _S. cyanosticta_ and _S. baudini_ are alike in certain +respects. The squamosals of both species are large and connected to the +maxillary by a bony connection; the squamosals of _S. phaeota_ are +large, but extend only two-thirds of the distance from the dorsal end of +the quadrate to the maxillary. In _S. baudini_ and _S. cyanosticta_ the +nasals are separated throughout their lengths from the ethmoid, whereas +the nasals of _S. phaeota_ are separated from the ethmoid by cartilage. +The latter separation is due to an incomplete ossification of the nasals +in _S. phaeota_. The bony part of each nasal is constricted in the +middle of the long axis of the bone, and the nasals are widest +anteriorly; posteriorly each nasal bears a medial process, which is +narrowly separated from the lateral edge of the ethmoid. + + + Table 5.--Variation in the Number of Teeth in the Species of + Smilisca. (All Are Males; N = Number of Jaws, or Twice the Number + of Individuals; Means Are Given in Parentheses After the Observed + Ranges.) + + =================+====+==============+==============+=========== + Species | N | Maxillary | Premaxillary | Vomerine + -----------------+----+--------------+--------------+----------- + _S. baudini_ | 20 | 49-65 (56.0) | 9-16 (13.6) | 5-9 (7.2) + _S. cyanosticta_ | 8 | 50-64 (57.9) | 10-12 (10.8) | 4-11 (7.1) + _S. phaeota_ | 20 | 50-68 (58.1) | 10-15 (12.1) | 5-9 (7.3) + _S. puma_ | 6 | 60-67 (63.6) | 11-13 (12.0) | 4-7 (5.3) + _S. sila_ | 8 | 48-60 (52.9) | 10-14 (11.3) | 5-7 (5.7) + _S. sordida_ | 12 | 39-55 (44.2) | 7-11 (9.3) | 4-6 (5.2) + -----------------+----+--------------+--------------+----------- + + +The teeth of all species of _Smilisca_ are spatulate and bifid. The +numbers of maxillary, premaxillary, and vomerine teeth are summarized in +Table 5. Smaller and presumably younger specimens of all species of +_Smilisca_ have fewer teeth than do larger specimens of the same +species. This correlation between size and number of teeth does not +exist as an interspecific trend within the genus; for example, the +smallest species in the genus, _S. puma_, has the highest number of +maxillary teeth. In small specimens of a given species wide gaps are +present between the maxillary teeth posteriorly; in large specimens the +gaps are filled by teeth, beginning anteriorly and progressing +posteriorly, until the maxillary dentition is continuous. + + +Musculature + +No extensive study of the muscular system was undertaken, but certain +muscles know to be of taxonomic importance were studied. + +_Jaw Musculature._--Starrett (1960) pointed out the unique jaw +musculature in _Smilisca_. In this genus M. depressor mandibulae +consists of two parts, one arising from the dorsal fascia and one from +the posterior arm of the squamosal. Two muscles arise from the anterior +arm of the squamosal and insert on the lateral face of the mandible. Of +these muscles, M. adductor mandibulae posterior subexternus lies medial +to the mandibular branch of the trigeminal nerve; the other, M. adductor +mandibulae externus superficialis, lies lateral to the same nerve (Fig. +9). In most other hylids the latter muscle is absent. No significant +variation in the position of the muscles was noted in the various +species of _Smilisca_, though M. adductor mandibulae originate somewhat +more anteriorly in _S. baudini_ and _S. cyanosticta_ than in the other +members of the genus, all of which have a shorter anterior arm of the +squamosal that does not reach the maxillary. The two separate parts of +M. depressor mandibulae are not so widely separated in members of the +_sordida_ group as in the _baudini_ group. + + [Illustration: FIG. 9. Lateral view of the left jaw of _Smilisca + baudini_; _A. M. E. S._, adductor mandibulae externus + superficialis; _A. M. P. S._, adductor mandibulae posterior + subexternus; _Col._, columella; _D. M._ depressor mandibulae; + _M. B. T. N._, mandibular branch trigeminal nerve; + _Sq._, squamosal. KU 64214, x5.] + + [Illustration: FIG. 10. Ventral view of throat musculature in an + adult male _Smilisca baudini_ (Superficial musculature on left, + deep musculature on right); _A. C._ anterior cornua of hyoid; + _Gen. L._, geniohyoideus lateralis; _Gen. M._, geniohyoideus + medialis; _Hyo._, hyoglossus; _Omo._, omosternum; _Pet._, + petrohyoideus; _S._, submentalis; _Sm._, submaxillaris; + _St._, sternohyoideus; _V. S._, vocal sac. KU 64220, x 2.5.] + +_Throat Musculature._--The frogs that comprise the genus _Smilisca_ are +characterized by paired subgular vocal sacs, essentially the same as +those in _Triprion_ (Duellman and Klaas, 1964). The following +description is based on _Smilisca baudini_ (Fig. 10). + +M. submentalis lies in the anterior angle of the lower jaw, is thick, +and consists of transverse fibers extending between the dentaries. M. +submaxillaris is thin and arises from the whole of the inner surface of +the lower jaw, except for the anterior angle occupied by M. submentalis. +Anteriorly M. submaxillaris is broadly attached by fascia to M. +hyoglossus and M. geniohyoideus, which lie dorsal to M. submaxillaris. +Medially this attachment continues posteriorly for about one-half the +length of the hyoglossus. Posteriorly M. submaxillaris is folded and +attached to M. sternoradialis of the pectoral girdle. The vocal sacs are +formed by a pair of posterolateral evaginations of M. submaxillaris; a +broad connection between the pouches allows free passage of air between +the pouches. + +The deeper throat musculature is essentially the same as that described +for _Phrynohyas spilomma_ by Duellman (1956), except for slight +differences in the place of attachment on the hyoid. + + +Skin + +_Structure_ + +The skin of _Smilisca_ is typical of that of most hylids in organization +and structure. _Smilisca sila_ is distinguished from other members of +the genus by the presence of small wartlike protrusions and peculiar +white, pustular spots on the dorsum. The wartlike structures are +composed of three or four epidermal cells, which protrude from the +surface of the epidermis; the structures are covered by a slightly +thickened layer of keratin. The white pustules are slightly elevated +above the surrounding skin. Internally they consist of aggregations of +swollen, granular, pigment-cells (perhaps lipophores) lying between the +epidermis and the melanophores. + +_Biochemical Variations_ + +Dried skins of all species of _Smilisca_ were sent to Jose M. Cei, +Instituto Nacional de Cuyo, Mendoza, Argentina, for biochemical +screening by means of the chromatographic techniques described by +Erspamer and Cei (1963). The species in the _baudini_ group have +detectable amounts of penta-hydroxi-trypatamine, whereas only a trace is +present in the other species. Furthermore, species in the _baudini_ +group differ from _S. sila_ and the _sordida_ group in lacking, or +having only a trace of, tryptophan-containing polypeptides. These +superficial biochemical tests support the arrangement of species as +ascertained by conventional taxonomic characters. + + +External Morphological Characters + +The features of external morphology that were studied in connection with +the taxonomy of the genus _Smilisca_ are discussed below. + +_Size and Proportions_ + +The frogs of the genus _Smilisca___ are medium to large tree frogs. The +three species comprising the _baudini_ group (_S. baudini_, +_cyanosticta_, and _phaeota_) are notably larger than _S. puma_, _sila_, +and _sordida_ (Table 6). The largest specimen that we examined is a +female of _S. baudini_ having a snout-vent length of 90 mm. _Smilisca +puma_ is the smallest species; the largest male has a snout-vent length +of 38 mm. and the largest female, 46 mm. + + + Table 6.--Comparison of Sizes and Certain Proportions of the Species + of Smilisca. (Means in Parentheses Below Observed Ranges; Data for + Males Only.) + + ================+====+===========+=============+===========+ + | | Snout-vent|Tibia length/| Tympanum/ | + Species | N | length | snout-vent | eye | + ----------------+----+-----------+-------------+-----------+ + | | | | | + _S. baudini_ |140 | 47.3-75.9 | 42.1-53.6 | 56.1-94.4 | + | | (58.7) | (47.8) | (73.5) | + | | | | | + _S. cyanosticta_| 40 | 44.6-56.8 | 51.9-59.7 | 62.7-88.4 | + | | (50.7) | (56.0) | (71.4) | + | | | | | + _S. phaeota_ | 50 | 40.8-65.5 | 50.9-60.2 | 62.7-85.5 | + | | (53.9) | (55.5) | (76.6) | + | | | | | + _S. puma_ | 20 | 31.9-38.1 | 48.2-53.1 | 52.1-72.2 | + | | (34.7) | (51.3) | (64.9) | + | | | | | + _S. sila_ | 33 | 31.6-44.8 | 49.7-58.1 | 47.6-58.3 | + | | (37.7) | (54.8) | (53.2) | + | | | | | + _S. sordida_ | 55 | 31.9-44.6 | 50.5-57.1 | 46.5-57.1 | + | | (37.9) | (53.4) | (49.1) | + ----------------+----+-----------+-------------+-----------+ + + +No outstanding differences in proportions exist between species, +although certain proportions are sufficiently different in some species +to warrant mention. _Smilisca baudini_ is a more squat and stocky frog +than other members of the genus; this is reflected in the somewhat +shorter hind legs (Table 6). The size of the tympanum relative to that +of the eye is highly variable within samples of a given species. Even +so, noticeable differences in the tympanum/eye ratio are apparent. +Members of the _baudini_ group have the largest tympani, whereas _S. +sila_ and _sordida_ have the smallest, and _S. puma_ is intermediate +(Table 6). + +_Shape of Snout_ + +Although all members of the genus have rather truncate snouts, subtle +differences exist among the species (Pl. 12). _Smilisca sila_ has the +shortest snout; that of _S. baudini_ is only slightly longer. The snouts +of _S. cyanosticta_ and _puma_ are nearly square in lateral profile, +whereas those of _S. phaeota_ and _sordida_ are slightly inclined. The +shape of the snout is relatively uniform within each species and +displays no noticeable sexual dimorphism, except in _S. sordida_, in +which there are sexual differences and geographic variation (see p. +324). + +_Hands and Feet_ + +The characters of the hands and feet are among the most taxonomically +important external features in _Smilisca_. Consistent differences exist +in relative lengths of the digits, size of subarticular tubercles, size +and number of supernumerary tubercles, size and shape of the inner +metatarsal tubercle, and amount of webbing (Pls. 4 and 5). In the +_baudini_ group the series of species (_baudini-phaeota-cyanosticta_) +show a progressive increase in amount of webbing in the hand and a +decrease in number, and corresponding increase in size, of supernumerary +tubercles. The amount of webbing in the feet of _S. baudini_ and +_phaeota_ is about the same, but the webbing is slightly more extensive +in _S. cyanosticta_. _Smilisca puma_ is unique in the genus in lacking +webbing in the hand; furthermore, this species is distinctive in having +many large subarticular tubercles on the hand and a relatively small +inner metatarsal tubercle. The two stream-inhabitants, _S. sila_ and +_sordida_, have shorter and stouter fingers than the other species. The +webbing is most extensive in both the hands and feet of these species, +which also are distinctive in having many small supernumerary tubercles +on the feet. + +_Ontogenetic Changes_ + +Minor ontogenetic changes in structure involve the shape of the snout, +relative size of the eye, development of the tympanum, and amount of +webbing in the hand. In recently metamorphosed young the snout is more +rounded than in adults; the canthus and loreal concavity are not +evident. Usually the tympanum is not differentiated in recently +metamorphosed young, and the eye is proportionately large. The webbing +in the feet is completely developed at metamorphosis, but young +individuals have noticeably less webbing in the hand than do adults of +the same species. + + +Coloration + +Some of the most distinctive characters of the species of _Smilisca_ are +color and pattern of the living frogs. Although many chromatic features +are lost or subdued in preserved specimens, the patterns usually +persist. + +_Metachrosis_ + +Change in color, well known in frogs, is common in hylids, especially in +species having green dorsal surfaces (_Phyllomedusa_ is a notable +exception). The non-green _Smilisca_ (_puma_, _sila_, and _sordida_) +changes color, but this mostly is a change in intensity of color. In +these species the markings usually are most distinct at night; +frequently by day the frogs become pallid. The most striking examples of +metachrosis in _Smilisca_ are found in the _baudini_ group, in which the +dorsal ground-color changes from green to tan; correlated with the +change in ground-color may be a corresponding change in the dorsal +markings, but the dorsal markings may change to the opposite color. + + +Chromosomes + +Chromosomes of all six species of _Smilisca_ were studied by means of +the propriono-orcein squash technique described by Duellman and Cole +(1965). Karyotype analysis was attempted for several species by means of +intraperitoneal injections of colchicine, which affected the mitotic +cells as desired, but the testes examined contained too few mitotic +cells to allow accurate determination of karyotypes. + +Haploid (_n_) chromosome numbers were determined from cells in +diakinesis, metaphase I, and metaphase II of meiosis. Diploid (2_n_) +chromosome numbers were determined from cells in late prophase and +metaphase of mitosis. Chromosome counts from as few as 23 meiotic cells +of _S. phaeota_ and as many as 80 cells of _S. sordida_ reveal a +constant haploid (_n_) number of 12; counts of chromosomes in one to +five mitotic cells in all species, except _S. sila_, reveal that the +diploid (2_n_) number is 24. + + + + +NATURAL HISTORY + + +Breeding + +Like most hylid frogs _Smilisca_ is most readily collected and observed +when individuals congregate for breeding. + +_Time of Breeding_ + +_Smilisca_ breeds primarily in quiet water and reaches its height of +breeding activity at times of plentiful rainfall,--usually from May +through October. Through most of its range _Smilisca baudini_ breeds in +those months, but in some places where abundant rain falls in other +seasons, the species breeds at those times. For example, in southern El +Peten and northern Alta Verapaz, Guatemala, _Smilisca baudini_ has been +found breeding in February and March. The other pond-breeding species +(_S. cyanosticta_, _phaeota_, and _puma_) live in regions lacking a +prolonged dry season, and possibly they breed throughout the year, but +breeding activity seems to be greatest in the rainiest months. + +The two stream-breeding species (_S. sila_ and _sordida_) breed in the +dry season when the streams are low and clear, principally in December +through April. At high elevations the species sometimes breed in the +rainy season; also, individuals sometimes breed in the short dry season +(summer canicula) in July and August. + +At several localities species have been found breeding at different +times of the year: _S. baudini_ in March and July at Chinaja, Guatemala; +_S. phaeota_ in April and August at Palmar Sur, Costa Rica; _S. puma_ in +February and July at Puerto Viejo, Costa Rica; and _S. sila_ in +February, April, and August at El Volcan, Panama. These observations +indicate only that the population breeds at more than one time in the +year, but do not provide any evidence on the breeding cycles of the +individual frogs. This is one important aspect of the natural history of +_Smilisca_ for which we lack data. + +_Breeding Sites_ + +All members of the genus _Smilisca_ presumably deposit their eggs in +water. + +_Smilisca baudini_ usually breeds in temporary rain pools; often these +are nothing more than shallow, muddy puddles. In other instances the +sites are extensive ditches or large flooded areas (Pl. 8, Fig. 1). This +species is an opportunistic breeder, and males gather at any of a wide +variety of suitable breeding sites that are formed by torrential rains +in the early part of the rainy season. _Smilisca baudini_ nearly always +breeds in open pools having bare earthen edges. Frequently congregations +of _S. baudini_ are found at such small pools, but are absent from +nearby large ponds surrounded by vegetation. + +Little is known of the breeding habits of _S. cyanosticta_, which +inhabits humid forests on foothills and lowlands. Apparently its +breeding sites are not unlike those of _S. phaeota_, which usually are +pools surrounded by vegetation (Pl. 8, Fig. 2), although sometimes males +of _S. cyanosticta_ call from open muddy puddles. In uplands, where +standing water is uncommon, this species breeds in quiet pools in +streams. + +_Smilisca puma_ breeds in grass-choked ponds and marshes, where the +males call from bases of dense clumps of grass in the water (Pl. 9, Fig. +1). + +_Smilisca sila_ and _S. sordida_ differ noticeably from other species in +the genus by breeding exclusively in streams, where males usually call +from rocks or gravel bars in or at the edges of streams (Pl. 9, Fig. 2); +sometimes individuals perch on bushes overhanging streams. In the +streams, or parts of streams, utilized by these frogs the water is +clear, shallow, and has a slow gradient; occasional males have been +found calling along cascading mountain streams. + +Breeding choruses composed of ten or more species of frogs are not +uncommon in Middle America, but _Smilisca_ usually breeds alone or with +one or two other species and at the most five others. This tendency +towards solitary breeding possibly is the result of selection of +breeding sites that are unsuitable to many other species of frogs. +Nevertheless, many other species of frogs have been found at the +breeding sites with the various species of _Smilisca_; these breeding +associates (Table 7) are most numerous for _S. baudini_, which has a +broad geographic range, including a variety of habitats. + +_Breeding Behavior_ + +_Calling sites._--All species of _Smilisca_ usually call from the +ground, including rocks and gravel bars; some individuals sit in shallow +water near the edge of the pool or stream. Sometimes males of _S. +baudini_, _sila_, and _sordida_ call from low bushes or trees near the +breeding site. One _S. baudini_ was observed calling while it was +floating on the surface of a pond. _Smilisca cyanosticta_, _phaeota_, +and _puma_ call from secluded places at the edge of the water or in the +water, whereas _S. baudini_, _sila_ and _sordida_ call from open +situations. + + + Table 7.--Breeding Associates of the Various Species of Smilisca. + + ==============================+========+============+========+=====+=====+======== + Associate |_S. |_S. |_S. |_S. |_S. |_S. + |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_ + ------------------------------+--------+------------+--------+-----+-----+-------- + _Rhinophrynus dorsalis_ | X | - | - | - | - | - + _Leptodactylus bolivianus_ | - | - | X | - | - | - + _Leptodactylus labialis_ | X | - | X | - | - | - + _Leptodactylus melanonotus_ | X | - | X | X | X | - + _Leptodactylus occidentalis_ | X | - | - | - | - | - + _Leptodactylus quadrivittatus_| - | - | X | - | - | - + _Leptodactylus pentadactylus_ | - | - | X | X | - | X + _Engystomops pustulosus_ | X | - | X | - | - | - + _Bufo canaliferus_ | X | - | - | - | - | - + _Bufo cavifrons_ | - | X | - | - | - | - + _Bufo coccifer_ | X | - | - | - | - | - + _Bufo coniferus_ | - | - | X | - | - | - + _Bufo cristatus_ | - | X | - | - | - | - + _Bufo gemmifer_ | X | - | - | - | - | - + _Bufo haematiticus_ | - | - | X | - | X | X + _Bufo kellogi_ | X | - | - | - | - | - + _Bufo luetkeni_ | X | - | - | - | - | - + _Bufo marinus_ | X | - | X | X | X | X + _Bufo marmoreus_ | X | - | - | - | - | - + _Bufo mazatlanensis_ | X | - | - | - | - | - + _Bufo melanochloris_ | - | - | X | - | X | X + _Bufo perplexus_ | X | - | - | - | - | - + _Bufo typhonius_ | - | - | X | - | X | - + _Atelopus varius_ | - | - | - | - | X | X + _Diaglena reticulata_ | X | - | - | - | - | - + _Diaglena spatulata_ | X | - | - | - | - | - + ------------------------------+--------+------------+--------+-----+-----+-------- + + Table 7.--_Continued_ + ==============================+========+============+========+=====+=====+======== + Associate |_S. |_S. |_S. |_S. |_S. |_S. + |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_ + ------------------------------+--------+------------+--------+-----+-----+-------- + _Hyla boulengeri_ | - | - | X | - | - | - + _Hyla colymba_ | - | - | - | - | X | - + _Hyla ebraccata_ | X | - | X | - | - | - + _Hyla elaeochroa_ | - | - | X | X | - | - + _Hyla eximia_ | X | - | - | - | - | - + _Hyla legleri_ | - | - | - | - | - | X + _Hyla microcephala_ | X | - | X | - | - | - + _Hyla phlebodes_ | - | - | X | X | - | - + _Hyla picta_ | X | - | - | - | - | - + _Hyla robertmertensi_ | X | - | - | - | - | - + _Hyla rosenbergi_ | - | - | X | - | - | - + _Hyla rufioculis_ | - | - | - | - | - | X + _Hyla smithi_ | X | - | - | - | - | - + _Hyla staufferi_ | X | - | - | - | - | - + _Hyla walkeri_ | X | - | - | - | - | - + _Phrynohyas inflata_ | X | - | - | - | - | - + _Phrynohyas spilomma_ | X | - | - | - | - | - + _Phrynohyas venulosa_ | X | - | - | - | - | - + _Phyllomedusa callidryas_ | X | - | X | - | - | - + _Phyllomedusa dacnicolor_ | X | - | - | - | - | - + _Phyllomedusa moreleti_ | X | X | - | - | - | - + _Pternohyla fodiens_ | X | - | - | - | - | - + _Smilisca baudini_ | X | X | - | - | - | - + _Smilisca cyanosticta_ | X | X | - | - | - | - + _Smilisca phaeota_ | - | - | X | - | - | - + _Smilisca puma_ | - | - | - | X | - | - + ------------------------------+--------+------------+--------+-----+-----+-------- + + Table 7.--_Concluded_ + ==============================+========+============+========+=====+=====+======== + Associate |_S. |_S. |_S. |_S. |_S. |_S. + |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_ + ------------------------------+--------+------------+--------+-----+-----+-------- + _Smilisca sila_ | - | - | - | - | X | X + _Smilisca sordida_ | - | - | X | - | X | X + _Triprion petasatus_ | X | - | - | - | - | - + _Cochranella fleischmanni_ | - | - | - | - | X | X + _Centrolene prosoblepon_ | - | - | - | - | X | - + _Gastrophryne elegans_ | X | - | - | - | - | - + _Gastrophryne olivacea_ | X | - | - | - | - | - + _Gastrophryne usta_ | X | - | - | - | - | - + _Hypopachus alboventer_ | X | - | - | - | - | - + _Hypopachus caprimimus_ | X | - | - | - | - | - + _Hypopachus inguinalis_ | X | - | - | - | - | - + _Hypopachus maculatus_ | X | - | - | - | - | - + _Hypopachus oxyrrhinus_ | X | - | - | - | - | - + _Hypopachus variolosus_ | X | - | - | - | - | - + _Rana palmipes_ | X | - | X | X | - | - + _Rana pipiens_ | X | - | - | - | - | - + _Rana warschewitschi_ | - | - | X | - | X | X + ------------------------------+--------+------------+--------+-----+-----+-------- + + +_Chorus structure._--Limited observations on some of the species of +_Smilisca_ show a definite organization of the calling behavior of +individuals. _Smilisca baudini_ and _S. phaeota_ call in duets. This is +especially noticeable in _S. baudini_, in which the members of a duet +often call from sites separated by only a few centimeters. The call of +_S. baudini_ consists of a series of like notes (see description of call +in following section); the duration of each note is about equal to the +interval between notes. Normally one individual utters one note, pauses, +and utters a single note again, or series of two or three notes. If +there is no response, the first individual often waits several seconds +or even several minutes and then repeats the call. The second individual +usually responds after the first or second note of the sequence. The +notes of the second individual usually are spaced so that they are +emitted in the intervals between the notes of the first individual. This +can be shown diagrammatically by having the figure "1" represent notes +of the first individual and figure "2," the notes of the second; an +empty interval is represented by "0": + + 1-0-1-2-1-2-1-2-1-2-1-2 + +Usually a chorus is initiated by one duet and is quickly picked up by +other individuals also calling in duets. A numerical representation of a +chorus of eight frogs would approximate the following organization: + + 1-0-1-2-1-2-1-2-1-2-1-2-1-2 + 3-0-3-4-3-4-3-4-3-4-3-4-3 + 5-6-5-6-5-6-5-6-5-6-5-6 + 7-8-7-8-7-8-7-8-7-8-7-8 + +After the first one or two duets are initiated, the second individuals +in the following duets usually call immediately after their respective +partners have given the first notes. The other noteworthy aspect about +the organization is that the entire chorus usually stops abruptly. +Normally the first duet stops calling shortly before the others, but +this is not invariable. Often one duet or one individual will emit +several notes after the rest of the frogs have become silent. An +interval of several minutes sometimes elapses before the chorus begins +again. Successive choruses apparently are initiated by the same duet. +Responses can be initiated artificially by imitating the call, and +sometimes any loud noise will start a chorus. + +Similar duets have been observed in _S. phaeota_. In this species the +intervals are often much longer than the notes, and if two males are +calling in close proximity, their calls can be mistaken for those of one +individual. _Smilisca phaeota_ does not congregate in large numbers; +usually only two males call from one restricted site. + +_Smilisca sila_ has a call consisting of a primary note followed by one +or more secondary notes. Males often call in duets, but not necessarily +so. In a duet, the first male usually utters only primary notes until +the second individual responds; then each individual produces a rapid +series of secondary notes. + +_Smilisca puma_ also produces primary and secondary notes. Although +individuals sometimes call alone, duets, trios, or quartets were more +common. The chorus is initiated by one individual uttering primary notes +until joined by the second, third, and fourth frogs. In one quartet in a +marsh 7.5 kilometers west of Puerto Viejo, Costa Rica, on February 19, +1965, the same individual initiated four consecutive choruses. Each time +the second member of the chorus was the same; the third and fourth frogs +joined the chorus nearly simultaneously. + +Individuals of _S. sordida_ are usually irregularly situated along a +stream. No duets or other combinations of individuals are apparent in +the chorus structure, but once an individual calls, a frog nearby calls +almost immediately; then a frog near the second individual calls, and so +on. The resulting series of calls gives the impression that the sound is +moving along the stream as successive individuals join the chorus and +the first callers become quiet. It is not known if the same individual +initiates successive choruses or if the order of calling is the same in +subsequent choruses. + +These limited observations on chorus structure in _Smilisca_ show the +presence of behavioral organization. The methods of establishing the +organization and the significance of the call-order in breeding have yet +to be discovered. + +Calling males of _S. baudini_ are often close together; some individuals +have been observed almost touching one another, but no indication of +territoriality or aggressive behavior has been witnessed. The more +distant spacing of the stream-breeding species _S. sila_ and _S. +sordida_ may be a function of calling-territories, but no direct +evidence is available to substantiate this supposition. + +_Sex recognition and amplexus._--Observations on _Smilisca baudini_ +indicate that the calls of males attract females. At Tehuantepec, +Oaxaca, Mexico, a female was first observed about two meters away from a +male calling at the edge of a rain pool; in a series of short hops she +progressed directly towards the male, although vegetation obscured him +until she was less than a meter away. When she approached to within +about 20 centimeters of the male, he took notice of her, moved to her, +and clasped her. At Chinaja, Alta Verapaz, Guatemala, a female swam +directly across a pool about three meters wide to a calling male. Her +line of movement took her within a few centimeters of a silent male, to +whom she paid no attention. She stopped just in front of the calling +male, which immediately clasped her. At a large muddy pond 4 kilometers +west-northwest of Esparta, Puntarenas, Costa Rica, a female was observed +swimming toward a small submerged tree; a male was calling from a branch +about one meter above the water. The female climbed to a branch about 20 +centimeters below the male, which upon seeing her there immediately +jumped down and clasped her. These few observations of _S. baudini_ show +that in this species females are capable of locating calling males by +means of phono-orientation; visual reception on the part of females +seems to be secondary. Contrariwise, males apparently become aware of +the proximity of females by seeing them; once a male sees a female he +usually tries to clasp her. Possibly the males receive stimuli by means +of chemo-reception, but in each observed instance the male obviously +looked at the female. + +Amplexus is axillary in all members of the genus. Normally amplexing +males hunch their backs and press their chins to the females' backs. +Clasping pairs are usually found at the edge of the water, but sometimes +amplexus takes place in trees or bushes. + +_Egg deposition._--Oviposition has been observed only in _Smilisca +baudini_. On the night of June 28, 1961, at Chinaja, Alta Verapaz, +Guatemala, a clasping pair was observed at the edge of a shallow rain +pool. After sitting for several minutes in shallow water, the female +(with male on her back) swam part way across the pool and grasped an +emergent stick with one hand. The female's body was nearly level with +the surface of the water, and her hind legs were outstretched as +deposition commenced; eggs were extruded rapidly. After a few seconds +the female moved slowly to another twig a few centimeters away and +deposited more eggs. This process was repeated until the female was +spent. The spawn resulted in a surface film covering roughly one square +meter. It is doubtful if this type of egg deposition occurs in any other +species in the genus, especially those that lay their eggs in streams. + +_Breeding Call_ + +The breeding calls of the six species of _Smilisca_ are alike in their +explosive nature. Calls are emitted quickly with a short burst of air +filling the vocal sac, which immediately deflates. Phonetically the +calls can be described as a single "wonk" or series of such notes in _S. +baudini_ and _S. cyanosticta_, a low growl in _S. phaeota_, a relatively +high pitched rattle in _S. sordida_, and a low squawk usually followed +by one or more rattling secondary notes in _S. puma_ and _S. sila_. +Quantitatively, the calls of the six species differ in number of notes, +duration of notes, and in pitch (Table 8, Pls. 10 and 11). Although no +measurements were taken on the intensity of the calls, we observed in +the field that each of the species has a loud voice. The call of _S. +baudini_ seems to carry farther than any of the others. + + + Table 8.--Comparison of Breeding Calls in Smilisca. (Observed Range + Given in Parentheses Below Mean. In Species Having Primary and + Secondary Notes, Only the Primary Notes Are Analyzed Here.) + + ==============+====+=======+===========+=========+=========+=======================+ + | | Notes | | | Funda- | Major | + Species | | per | Duration | Pulses | mental | frequencies (cps) | + | N | call | of note | per |frequency+-----------+-----------+ + | | group | (seconds) | second | (cps) | Lower | Upper | + --------------+----+-------+-----------+---------+---------+-----------+-----------+ + _S. | 20 | 8.0 | 0.11 | 174.7 | 166.2 | 351 | 2507 | + baudini_ | | (2-15)|(0.09-0.13)|(140-195)|(135-190)| (175-495) |(2400-2725)| + | | | | | | | | + _S. | 10 | 1.2 | 0.38 | 147.0 | 145.1 | 841 | 1894 | + cyanosticta_| | (1-2) |(0.25-0.45)|(110-180)|(135-160)| (480-975) |(1600-2100)| + | | | | | | | | + _S. phaeota_ | 10 | 1.6 | 0.31 | 116.0 | 143.0 | 372 | -- | + | | (1-2) |(0.10-0.45)|(100-130)|(110-165)| (330-495) | -- | + | | | | | | | | + _S. puma_ | 28 | 3.7 | 0.13 | 208.2 | 145.6 | 743 | 1868 | + | | (2-10)|(0.06-0.35)|(187-240)|(125-200)| (495-980) |(1456-2240)| + | | | | | | | | + _S. sila_ | 15 | 2.4 | 0.16 | 108.5 | 103.0 | 899 | 2218 | + | | (1-6) |(0.06-0.28)| (97-120)| (90-115)| (665-1180)|(1980-2700)| + | | | | | | | | + _S. sordida_ | 19 | 1.7 | 0.29 | 104.7 | 123.1 | 1216 | 2694 | + | | (1-6) |(0.18-0.45)| (78-135)| (90-140)|(1150-1540)|(2340-2990)| + --------------+----+-------+-----------+---------+---------+-----------+-----------+ + + +_Call rate._--The rate at which call-groups are produced varies from one +every few seconds to one in several minutes. In _S. baudini_, +_cyanosticta_, _phaeota_, and _sordida_, call-groups are produced as +frequently as every 12 seconds, but usually more time elapses between +call groups. In _S. sordida_, five or more minutes sometimes elapse +between call-groups. The interval is somewhat less in _S. phaeota_. +Calls are repeated at much shorter intervals in _S. puma_ (5-55 seconds) +and _S. sila_ (4-20 seconds). + +_Notes per call-group._--Except for _S. puma_ and _S. sila_, the series +of notes produced in any given call of a species of _Smilisca_ is +essentially the same; there is no differentiation into primary and +secondary notes. _Smilisca cyanosticta_ and _S. phaeota_ emit only one +or two relatively long notes per call-group, whereas _S. baudini_ and +_S. sordida_ produce as many as 15 and 6 notes, respectively. Males of +_S. puma_ and _S. sila_ often produce only the primary note; sometimes +this is done several times before the secondary notes are produced. For +example, one _S. puma_ (KU 91711; tape No. 379) produced the following +number of notes in consecutive call-groups: 1, 1, 1, 1, 2, 2, 3, 1, 4; +secondary notes are present in only four of the nine call-groups. A +typical series of consecutive call-groups in _S. sila_ (KU 91852; Tape +No. 385) has 1, 1, 1, 2, 4, 2 notes per call-group; secondary notes are +present in only half of the call-groups. _Smilisca puma_ apparently +always produces at least two primary notes before emitting secondary +notes; sometimes only primary notes are produced in one series of calls. +The number of secondary notes following a given primary varies from one +to nine; the modal number is one, and the mean is three in 27 +call-groups. _Smilisca sila_ frequently begins a series of calls with +two or more primary notes, but sometimes the first primary note is +followed immediately by two or more secondary notes. The number of +secondary notes following a given primary varies from one to five; the +modal number is one, and the average is two in 13 call-groups. + +_Duration._--The average duration of call-groups consisting of two or +more notes is 1.18 seconds in _S. baudini_; 1.02 in _cyanosticta_, 0.91 +in _phaeota_, 1.32 in _puma_, 1.48 in _sila_, and 1.29 in _sordida_. +Although there is considerable variation in the lengths of the notes +(only primary notes in _S. puma_ and _sila_ are considered here), _S. +cyanosticta_, _phaeota_, and _sordida_ have noticeably longer notes than +do the other species (Table 8). The secondary notes are longer than the +primary notes in _S. puma_ (average 0.27 secs. as compared with 0.13 +secs.) and in _S. sila_ (average 0.25 secs., as compared with 0.16 +secs.). + +_Note repetition rate._--The rate at which notes in call-groups +containing two or more notes are produced varies in _S. baudini_ from +2.5 to 7.1 (average, 3.7) calls per second; _cyanosticta_, 1.8-2.1 +(1.9); _phaeota_, 2.0-2.4 (2.2); _puma_, 1.9-2.9 (2.2); _sila_, 1.3-2.4 +(1.8); and _sordida_, 1.5-2.6 (2.1). _Smilisca baudini_, which has +notes of short duration (0.09 to 0.13 seconds), has the fastest +note-repetition rate. Although the individual notes of _S. cyanosticta_ +and _S. phaeota_ are relatively long (average, 0.38 and 0.31 seconds, +respectively), the intervals between the notes is short; consequently, +their note-repetition rates do not differ greatly from those of _S. +puma_ and _S. sila_, which have shorter notes (average, 0.13 and 0.16 +seconds, respectively) but longer intervals between notes. + +_Pulse rate._--Pulses vary in frequency from 78 to 240 per second in the +calls analyzed (only primary notes in _S. puma_ and _S. sila_), but the +variation in any given species is much less than that in the entire +genus (Table 8). _Smilisca puma_ is outstanding in having a high pulse +rate, which is approached only by that of _S. baudini_. Even in the +species having the lowest pulse rates, the pulsations are not audible. +The secondary notes produced by _S. puma_ and _S. sila_ have a slower +pulse rate than the primary notes; often the pulses are audible. In _S. +puma_ the pulse rate of secondary notes is sometimes as low as 48 pulses +per second, and in _S. sila_ still lower (as low as 40 pulses per +second). The upper limits of pulse rate in the secondary notes in these +species merge imperceptibly with the rates of the primary note; +consequently, on the basis of pulse rate alone it is not always possible +to distinguish primary from secondary notes. + +_Frequency._--_Smilisca_ produces noisy (as opposed to more musical) +calls, and the energy is distributed throughout the frequency spectrum; +the calls are poorly modulated, except in _S. sordida_, in which two +usually discrete bands of frequency are present (Pl. 11C). For the most +part the calls of _Smilisca_ consist of little modified energy of the +fundamental frequency and of its harmonics, some of which are +emphasized. + +The upper frequency range varies within each species and even within the +calls of one individual. _Smilisca phaeota_ has the lowest upper +frequencies; no calls ranged above 4400 cycles per second (cps.), and +half of the calls never exceeded 3000 cps. _Smilisca cyanosticta_ +produces calls in which the upper frequency is below 7000 cps. and +usually below 6000 cps. Likewise, _S. puma_ produces calls that are +below 7000 cps., whereas _S. sila_ has frequencies of up to 8400 cps. In +both _S. baudini_ and _S. sordida_, the highest frequencies attained are +about 9100 cps. Variation in the highest frequencies in a series of +consecutive calls by one individual frog was noted in all species. Such +variation is especially prevalent in _S. puma_; for example one +individual (KU 87771; Tape No. 376) recorded at a temperature of 24 deg. C. +at 7.5 kilometers west of Puerto Viejo, Heredia Province, Costa Rica, on +July 31, 1964, produced three consecutive primary notes having upper +frequencies of about 6000, 4000, and 4000 cps., respectively. Apparently +in a given species the production of the higher frequencies in some +notes and not in others is correlated with the amount of distention of +the vocal sac and is not dependent upon the structure or tension of the +vocal cords. + +Although the dominant frequency in _S. sordida_ is lower than that in +_S. baudini_ and _S. cyanosticta_, the call of the former is audibly +higher-pitched. This is due primarily to the emphasis on certain +harmonics at a high frequency (sometimes as high as 9000 cps.) in _S. +sordida_, whereas in _S. baudini_ and other species, if harmonics are +present at those frequencies, they are not emphasized. + +The fundamental frequencies are as low as 90 cps. in _S. sila_ and _S. +sordida_ and as high as 200 cps. in _S. puma_ (Table 8). The fundamental +frequency seemingly is relatively unimportant in determining the general +pitch of the call, a characteristic most dependent on the dominant +frequency and emphasized harmonics in the higher-frequency spectrum. In +none of the species is the fundamental the dominant frequency. In the +low-pitched call of _S. phaeota_ the dominant frequency is the third +harmonic (the second harmonic above the fundamental frequency, which is +the first harmonic). In all other species a much higher harmonic is +dominant; for examples, in _S. cyanosticta_ harmonics from 10 to 15 are +dominant; in _S. baudini_, 15-19; and _S. sila_, 20-30. + +A glance at the audiospectrographs and their accompanying sections (Pls. +10 and 11) reveals the presence of two emphasized bands of frequency in +all species except _S. phaeota_, in which only the lower band is +present. These two bands of emphasized harmonics are part of a +continuous, or nearly continuous, spread of energy throughout the +frequency spectrum, except in _S. sordida_ in which the bands are +usually distinct. As shown in the sections, certain harmonics in each of +the bands are emphasized with nearly equal intensity. Therefore, with +the exception of _S. phaeota_, the calls of _Smilisca_ are characterized +by two major frequencies, one of which is the dominant frequency and the +other is a subdominant frequency (Table 8). The upper major frequency is +dominant in all calls in _S. baudini_ and _S. cyanosticta_, but either +major frequency may be dominant in other species. The upper major +frequency is dominant in 65 per cent of calls by _S. puma_, 87 per cent +in _S. sila_, and 68 per cent in _S. sordida_. Individuals of these +three species sometimes produce a series of calls in which the dominant +frequency changes from one of the major frequencies to the other. Four +consecutive notes emitted by an individual of _S. sordida_ recorded 13 +kilometers east-northeast of Golfito, Puntarenas Province, Costa Rica, +had dominant frequencies of 910, 1950, and 750 cps., respectively. In +each case, an alternation of major frequencies took place in respect to +dominance. An individual of _S. puma_ from 7.5 kilometers west of Puerto +Viejo, Costa Rica, produced a primary note followed by one secondary +note; each note had major frequencies at 600 and 1800 cps.; the dominant +frequency of the primary note was at 1800 cps., whereas in the secondary +note the dominant frequency was at 600 cps. The difference in emphasis +on the major frequencies is so slight that shift in dominance is not +audible. + +_Effect of temperature on calls._--The present data are insufficient to +test statistically the correlation between temperature and variation +within certain components of the calls in _Smilisca_, but even a crude +graph shows some general correlations. The widest range of temperatures +is associated with the recordings of _S. baudini_. Three individuals +recorded at a temperature of 30 deg. C. at Tehuantepec, Oaxaca, had pulse +rates of 180 pulses per second and fundamental frequencies of 160-180 +cps., as compared with an individual recorded at a temperature of 17 deg. +C., which had a pulse rate of 140 and a fundamental frequency of 135 +cps. All individuals of _S. baudini_ recorded at higher temperatures had +faster pulse rates and higher fundamental frequencies. Pulse rates +differ in the other species in the genus but less strikingly (probably +owing to narrower ranges of temperatures at which recordings were made). +In five recordings of _S. sordida_ made at 20 deg. C. the pulse rate is +80-90, as compared with four recordings made at 25 deg. C. having pulse +rates of 120-135. Thirteen recordings of _S. sila_ made at 17 deg. C. have +pulse rates of 97-112 (average 105); one individual recorded at 26 deg. C. +has 120 pulses per second. Seemingly no correlation exists between +temperature and other characteristics of the calls, such as duration and +rate of note-repetition. + +_The breeding call as an isolating mechanism._--Blair (1958), Bogert +(1960), Duellman (1963a), Fouquette (1960), Johnson (1959), and others +have provided evidence that the breeding calls of male hylids (and +other anurans) serve as isolating mechanisms in sympatric species. In +summarizing this discussion of the breeding calls of _Smilisca_ we want +to point out what seem to be important differences in the calls that may +prevent interspecific hybridization in sympatric species of _Smilisca_. + +The genus is readily divided into two species-groups on morphological +characters; this division is supported by the breeding calls. In the +species of the _baudini_ group the calls are unmodulated and lack +secondary notes. In the _sordida_ group the calls either have secondary +notes or are modulated. + +_Smilisca baudini_ occurs sympatrically with _S. cyanosticta_ and _S. +phaeota_; where they occur together, both species sometimes breed in +like places at the same time. We are not aware of these species breeding +synchronously at exactly the same site, although _S. baudini_ and _S. +cyanosticta_ were calling on the same nights and less than 100 meters +apart in Oaxaca in June, 1964. Regardless of their respective breeding +habits, sympatric species have calls that differ notably. Except for the +higher fundamental and dominant frequencies, the calls of _S. +cyanosticta_ and _S. phaeota_ closely resemble one another, but the +calls of both species differ markedly from that of _S. baudini_. The +geographic ranges of _S. cyanosticta_ and _S. phaeota_ are widely +separated. + +The calls of the allopatric species _S. puma_ and _S. sila_ are not +greatly different. _Smilisca sordida_ has a distinctive call and occurs +sympatrically with _S. puma_ and _S. sila_. In the streams in southern +Costa Rica _S. sordida_ and _S. sila_ breed synchronously, but the +high-pitched modulated call of the former is notably different from the +lower, unmodulated call of _S. sila_. + +The data indicate that the calls of related sympatric species differ +more than the calls of related allopatric species. We postulate that +these differences evolved to support the reproductive isolation of the +sympatric species. The data are insufficient to determine geographic +variation in the calls and to determine if differences in the calls are +enhanced in areas of sympatry as compared with the allopatric parts of +the ranges. + +_Other calls._--As stated previously, there is no direct evidence of +territoriality in _Smilisca_; we have heard no calls that can be +definitely identified as territorial. Single notes of _S. baudini_, +_phaeota_, and _sila_ have been heard by day, just prior to rains, or +during, or immediately after rains. Such calls can be interpreted as +"rain calls," which are well known in _Hyla eximia_ and _Hyla +squirella_. Distress calls are known in several species of _Rana_ and in +_Leptodactylus pentadactylus_; such calls result from the rapid +expulsion of air over the vocal cords and with the mouth open. Distress +calls have been heard from _S. baudini_. At Charapendo, Michoacan, +Mexico, a male that had one hind limb engulfed by a _Leptodeira +maculata_ emitted several long, high-pitched cries. A clasping pair of +_S. baudini_ was found in a bush at the edge of a marshy stream 2 +kilometers northeast of Las Canas, Guanacaste Province, Costa Rica. When +the pair was grasped, the female emitted a distress call. + + +Eggs + +Eggs of _S. baudini_, _cyanosticta_, and _phaeota_ have been found in +the field, and eggs of _S. sila_ have been observed in the laboratory. +The eggs of _S. puma_ and _sordida_ are unknown. Insofar as known, +_Smilisca baudini_ is unique in the genus in depositing the eggs in a +surface film. Each egg is encased in a vitelline membrane, but +individual outer envelopes are lacking. The eggs are small; the diameter +of recently-deposited eggs is about 1.3 mm. and that of the vitelline +membrane is about 1.5 mm. The eggs of _S. cyanosticta_ and _phaeota_ are +deposited in clumps, and the eggs are larger than those of _S. baudini_. +Diameters of eggs of _S. cyanosticta_ are about 2.3 mm., and those of +the outer envelopes are about 4.0 mm. Artificially fertilized eggs of +_S. sila_ raised in the laboratory have diameters of about 2.4 mm.; the +diameter of the outer envelopes is about 4.9 mm. + +In order to determine the reproductive potential of the six species, +ovulated eggs were removed from females and counted. The numbers of eggs +recorded are: 3 _S. baudini_--2620, 2940, 3320; 1 _S. cyanosticta_--910; +3 _S. phaeota_--1665, 1870, 2010; 1 _S. puma_--518; 3 _S. sila_--369, +390, 473; 3 _S. sordida_--524, 702, 856. These limited data indicate +that the large species (_S. baudini_, _cyanosticta_, and _phaeota_) have +more eggs than do the smaller species. The stream-breeding species (_S. +sila_ and _sordida_) have relatively few eggs by comparison with the +pond-breeders. Possibly this is a function of size of eggs rather than a +correlation with the site of egg-deposition. + + +Tadpoles + +The acquisition of tadpoles of all of the species of _Smilisca_ has made +possible the use of larval characters in erecting a classification and +in estimating the phylogenetic relations of the several species. +Furthermore, developmental series of tadpoles of four species allow a +comparison of the growth and development in these species. Throughout +the discussion of tadpoles we have referred to the various developmental +stages by the Stage Numbers proposed by Gosner (1960). + +_General Structure_ + +Tadpoles of the genus _Smilisca_ are of a generalized hylid type, having +2/3 tooth-rows, unspecialized beaks, mouth partly or completely bordered +by papillae, lateral fold present in the lips, spiracle sinistral, anal +tube dextral, and caudal musculature extending nearly to tip of caudal +fin. Although minor differences exist in coloration, proportions, and +mouthparts, no great modifications of the basic structure are present. + +_Comparison of Species_ + +The larval characters of the species of _Smilisca_ are compared below +and illustrated in Figures 11-15. + +_Shape and Proportions._--The bodies of _S. baudini_, _cyanosticta_, +_phaeota_, and _puma_ are rounded and about as wide as deep; the eyes +are moderately large and directed dorsolaterally, and the nostrils are +about midway between the bluntly rounded snout and the eyes. The mouths +are medium-sized and directed anteroventrally. The bodies of tadpoles of +_S. sila_ and _sordida_ are slightly compressed dorso-ventrally. The +snout is moderately long and sloping; the eyes are larger and directed +more dorsally than in the other species, and the nostrils are closer to +the eyes than the snout. The mouths are moderately large and directed +ventrally. + +The tail is about half again as long as the body in _S. baudini_, +_cyanosticta_, _phaeota_, and _puma_; in these species the caudal +musculature is moderately heavy, and the caudal fins are deep. The +caudal musculature is upturned distally in _S. baudini_ and _phaeota_, +and the dorsal fin extends anteriorly onto the body in these two species +and in _S. puma_. The tail is about twice as long as the body in _S. +sila_ and _sordida_. In both species the caudal fins are shallow in +comparison with the depth of the caudal musculature, especially in _S. +sordida_ (Fig. 14); in neither species does the dorsal fin extend +anteriorly onto the body. + + [Illustration: FIG. 11. Tadpoles of _Smilisca baudini_: (A) Stage + 21 (KU 62155) x 10; (B) Stage 25 (KU 68467) x 5; (C) Stage 30 + (KU 60018) x 4; (D) Stage 41 (KU 60018) x 3.] + +_Mouthparts._--The mouth of _S. sordida_ is completely bordered by two +rows of papillae, whereas in the other species the median part of the +upper lip is devoid of papillae. _Smilisca baudini_ and _puma_ have two +rows of papillae; _S. sila_ has one complete row (except medially on the +upper lip) and one incomplete row, and _S. cyanosticta_ and _phaeota_ +have only one row (Fig. 15). All species have numerous papillae in the +lateral fold; the fewest lateral papillae are found in _S. cyanosticta_ +and _phaeota_. Although all species have two rows of teeth in the upper +jaw and three rows in the lower jaw, specific differences in the nature +of the rows exist between certain species. The second upper tooth-row is +narrowly interrupted medially in _S. sila_ and _sordida_ and broadly +interrupted in the other species. The first upper row is strongly arched +in _S. puma_, moderately arched in _S. baudini_ and _sila_, and weakly +arched in the other species. In all species the third lower tooth-row is +the shortest, only slightly so in _S. sila_ and _sordida_, but only +about half the length of the second lower row in _S. puma_. + + [Illustration: FIG. 12. Tadpoles of _Smilisca cyanosticta_: + (A) Stage 21 (KU 87648) (B) Stage 25 (KU 87651) x 5; + (C) Stage 30 (KU 87652) x 4; (D) Stage 40 (KU 87650) x 3.] + +The beaks are well developed and finely serrate in all species. The +lower, broadly V-shaped, beak is slender in _S. puma_, rather robust in +_S. baudini_ and _sila_, and moderately heavy in the other species. The +lateral processes of the upper beak are shortest in _S. puma_ and +longest in _S. baudini_ and _sordida_. In the latter the inner margin of +the upper beak and lateral process have the form of a shallow S, whereas +in the other species the inner margin of the upper beak forms a +continuous arch with the lateral processes (Fig. 15). + + [Illustration: FIG. 13. Tadpoles of _Smilisca phaeota_: (A) Stage 21 + (KU 68479) x 14; (B) Stage 25 (KU 68480) x 5; (C) Stage 30 (KU + 68482) x 4; (D) Stage 40 (KU 68483) x 3.] + +_Coloration._--The tadpoles of _Smilisca_ lack the bright colors or bold +markings characteristic of some hylid tadpoles; even so, the subdued +colors and arrangement of pigments provide some distinctive markings by +which the species can be distinguished from one another. The species +comprising the _baudini_ group (_S. baudini_, _cyanosticta_, and +_phaeota_) are alike in having the body brown or grayish brown dorsally +and transparent with scattered brown pigment ventrally. A cream-colored, +crescent-shaped mark is present on the posterior edge of the body; this +mark is usually most noticeable in _S. baudini_ and least so in _S. +cyanosticta_. Other differences in coloration in members of the +_baudini_ group are relative and subtle. _Smilisca phaeota_ usually is +more pallid than _baudini_, and _cyanosticta_ usually is darker than +_baudini_; both species have larger dark markings on the tail than does +_S. phaeota_. _Smilisca baudini_ has a dark streak on the middle of the +anterior one-fourth of the tail (Figs. 11-13). + +_Smilisca puma_ is distinctive in having a grayish brown body and dark +gray reticulations on the tail. _Smilisca sila_ and _sordida_ are +distinctive in having pairs (sometimes interconnected) of dark marks on +the dorsal surfaces of the caudal musculature, and in dorsal view the +tail appears to be marked with dark and pale creamy tan transverse bars. +These dark marks, as well as the small flecks on the tail, are brown in +_S. sila_ and red in _sordida_. _Smilisca sila_ has dark brown flecks on +the dorsal surface of the body and small white flecks laterally; these +markings are absent in _S. sordida_ (Fig. 14). + +Descriptions of the coloration of living tadpoles are given in the +accounts of the species. + + [Illustration: FIG. 14. Tadpoles of _Smilisca_; (A) _S. puma_, + Stage 30 (KU 91807); (B) _S. sila_, Stage 25 (KU 80260); + _S. sordida_, Stage 30 (KU 68475). All x 3.5.] + +_Growth and Development_ + +Information on the growth and development of Middle American hylids is +scanty. Adequate descriptions have been published for _Phyllomedusa +annae_ (Duellman, 1963b), _Phrynohyas venulosa_ (Zweifel, 1964), and +_Triprion petasatus_ (Duellman and Klaas, 1964). Material is available +for adequate descriptions of the developmental stages of four species of +_Smilisca_ (Tables 9-12, Figs. 11-13). Because none of the tadpoles was +raised from hatching to metamorphosis, the rate of growth and duration +of the larval stages are unknown. + + [Illustration: FIG. 15. Mouthparts of tadpoles of _Smilisca_; + (A) _S. baudini_ (KU 60018); (B) _S. puma_ (KU 91807); + (C) _S. cyanosticta_ (KU 87625); (D) _S. sila_ (KU 80620); + (E) _S. phaeota_ (KU 68482); (F) _S. sordida_ (KU 68475). + All x17.] + + Table 9.--Growth and Development of Tadpoles of Smilisca baudini. + (Means Are Given in Parentheses After the Observed Ranges.) + + ==================================================================== + Stage | N | Total length | Body length | Tail length + -------------------------------------------------------------------- + 21 |10 | 5.1-5.4 (5.22) | 2.6-2.7 (2.54) | 2.5-2.7 (2.58) + 24 |10 | 6.0-6.5 (6.20) | 2.3-2.6 (2.45) | 3.5-3.9 (3.69) + 25 |10 | 7.2-8.3 (7.78) | 3.0-3.3 (3.14) | 4.2-5.0 (4.64) + 27 |10 | 18.5-21.5 (20.22) | 8.0-9.0 (8.38) |10.4-13.0 (11.84) + 29 |10 | 21.5-24.5 (22.60) | 8.5-10.0 (9.25) |12.5-14.5 (13.35) + 37 | 3 | 28.5-31.0 (30.00) |11.0-12.5 (11.67) |17.5-19.0 (18.00) + 38 |10 | 35.0-37.5 (35.50) |12.0-13.5 (12.80) |21.5-24.0 (22.70) + 40 | 2 | 34.0-37.0 (35.50) |12.5-13.5 (13.00) |21.5-23.5 (22.50) + 41 |10 | 34.0-37.0 (35.50) |12.5-13.5 (13.00) |21.5-23.5 (22.50) + 42 | 3 | 24.0-30.0 (27.00) |12.5-13.0 (12.67) |11.5-17.0 (14.33) + 45 | 6 | 14.0-24.0 (17.58) |12.5-14.0 (13.37) | 1.5-10.0 (4.17) + 46 |23 | ---- |12.0-15.5 (13.34) | ---- + -------------------------------------------------------------------- + + + Table 10.--Growth and Development of Tadpoles of Smilisca + cyanosticta. (Means Are Given in Parentheses After the + Observed Ranges.) + + ====================================================================== + Stage | N | Total length | Body length | Tail length + ---------------------------------------------------------------------- + 21 | 10| 5.8-6.5 (6.28) | 2.8-3.1 (3.00) | 3.0-3.5 (3.28) + 25 | 10| 7.9-9.2 (8.44) | 2.7-3.2 (2.96) | 4.8-6.0 (5.48) + 30 | 7| 22.5-25.0 (23.50) | 8.5-9.5 (9.00) |14.0-15.5 (14.57) + 36 | 10| 27.0-30.0 (28.75) | 9.5-11.5 (10.80) |17.0-18.5 (17.95) + 42 | 2| 26.0-27.0 (26.50) | 10.00 |16.0-17.0 (16.50) + 46 | 2| -- | 14.00 | -- + ---------------------------------------------------------------------- + + +Hatchlings of three species (_S. baudini_, _cyanosticta_, and _phaeota_) +are available. These larvae have non-functional eyes and large oral +suckers. By the time the larvae have developed to stage 21, external +gills are present, the caudal musculature and caudal fin have been +differentiated, and the head is distinguishable from the body. In stage +21 oral suckers and a large amount of yolk are still present. + +The developmental data on the four species show no significant +variations; consequently, we will describe the development of only one +species, _Smilisca phaeota_ (Table 11, Figs. 13 and 16). + +_Stage 21._--Bulging cream-colored yolk mass, transparent cornea, and +moderately long, unbranched filamentous gills, and oral suckers present; +mouth having irregular papillae on lower lip; teeth and beaks absent; +caudal myomeres distinct; pigmentation uniform over body and caudal +musculature; caudal fin transparent with scattered small flecks. + + [Illustration: FIG. 16. Relative rate of growth in tadpoles of + _Smilisca phaeota_ as correlated with developmental stages. + Formulas for the limb bud refer to its length (L) in relation to + basal diameter (D).] + +_Stage 25._--Operculum complete; gills absent; sinistral spiracle +apparently functional; cloacal tail-piece, nasal capsules, and external +nares present; gut partly formed; mouth bordered by single row of +papillae, except medially; small papillae present in lateral fold of +lips; two upper and three lower tooth-rows present, but not fully +developed; beaks apparently fully developed; depth of dorsal and ventral +fins less than depth of caudal musculature: tip of tail upturned; +pigment on body most dense on dorsum and sides; faint, nearly +pigmentless crescent-shaped mark on posterior edge of body; +concentrations of pigment forming small spots on tail. + +_Stage 28._--Mouthparts complete; limb bud about half as long as thick; +other structural features and coloration closely resemble those in stage +25. + +_Stage 30._--Limb bud approximately twice as long as thick; body as deep +as wide; dorsal fin deepest just posterior to body; ventral fin deeper +than caudal musculature; tail sharply upturned distally; anal tube +dextral; brown pigment sparse on flanks. + + + Table 11.--Growth and Development of Tadpoles of Smilisca phaeota. + (Means Are Given in Parentheses After the Observed Ranges.) + + ==================================================================== + Stage | N | Total length | Body length | Tail length + -------------------------------------------------------------------- + 15 | 10| -- | 1.9-2.1 (1.97) | -- + 16 | 8| -- | 2.0-2.2 (2.07) | -- + 18 | 4| -- | 2.2-2.6 (2.31) | -- + 21 | 3| 7.9-8.6 (8.21) | 4.1-4.5 (4.31) | 3.8-4.1 (3.92) + 25 | 10| 8.7-10.6 (9.69) | 4.5-4.8 (4.64) | 4.3-6.0 (5.05) + 26 | 11| 12.3-16.1 (14.01) | 4.2-6.3 (5.60) | 6.7-9.8 (8.41) + 27 | 10| 13.0-15.7 (14.28) | 4.9-6.2 (5.40) | 7.7-10.5 (8.88) + 28 | 13| 13.9-20.9 (15.62) | 5.2-8.3 (5.75) | 8.5-12.6 (9.85) + 29 | 8| 17.8-22.3 (19.79) | 6.3-8.4 (7.19) | 11.5-14.0 (12.60) + 30 | 9| 20.3-24.8 (22.85) | 8.1-10.5 (9.32) | 10.5-15.5 (13.53) + 31 | 5| 24.1-28.5 (26.61) | 9.4-11.2 (10.59) | 14.7-17.3 (16.02) + 34 | 5| 24.8-29.4 (27.31) | 9.2-11.6 (10.73) | 15.6-18.5 (16.80) + 36 | 3| 30.0-30.1 (30.07) |10.1-12.2 (11.15) | 18.9-20.0 (19.44) + 37 | 4| 28.9-34.1 (31.75) |11.5-12.4 (11.88) | 17.4-22.5 (19.88) + 38 | 1| 28.98 |12.88 | 16.10 + 39 | 2| 35.6-36.9 (36.25) |14.00 | 21.6-22.9 (22.25) + 40 | 2| 32.3-39.8 (36.05) |14.00 | 18.3-21.8 (20.05) + 43 | 2| 21.5-23.0 (22.25) |14.2-14.8 (14.45) | 6.8-8.8 (7.80) + 44 | 4| -- |14.5-15.6 (15.08) | -- + 46 | 11| -- |12.7-16.7 (14.26) | -- + -------------------------------------------------------------------- + + + Table 12.--Growth and Development of Tadpoles of Smilisca sordida. + (Means Are Given in Parentheses After the Observed Ranges.) + + =======+===+==================+==================+================= + Stage | N | Total length | Body length | Tail length + -------+---+------------------+------------------+----------------- + 25 | 6 | 25.5-28.0 (26.1) | 9.0-9.5 (9.3) | 16.2-18.5 (16.7) + 33 | 2 | 28.5-30.0 (29.3) | 10.2-10.5 (10.4) | 18.0-19.8 (18.9) + 36 | 8 | 29.5-34.5 (32.3) | 10.2-11.7 (10.8) | 19.3-23.0 (21.5) + 37 | 7 | 31.6-37.5 (34.6) | 11.0-12.5 (11.5) | 21.6-25.0 (23.2) + 41 | 3 | 33.0-37.2 (35.2) | 11.6-12.2 (11.9) | 21.4-25.2 (23.2) + 43 | 1 | ---- | 12.4 | ---- + 46 | 9 | ---- | 13.1-15.7 (14.9) | ---- + -------+---+------------------+------------------+----------------- + + +_Stages 34_, _36_, _37_, and _38_.--Stage 34, foot paddle-shaped with +four toe buds; stage 36, five toe buds; stages 37 and 38, lengthening of +toes. In all four stages, spiracle persistent, and pigmentation +resembling that of early stages. + +_Stage 39._--Metatarsal tubercle present; greatest total length (36.9 +mm.) attained. + +_Stage 40._--Subarticular tubercles prominent; skin over forelimbs +transparent; cloacal tail-piece and spiracle absent; outer tooth-rows +degenerating; caudal fins shallower than in preceding stages; distal +part of tail nearly straight; size of dark markings on tail decreased; +pigment present on hind limb. + +_Stage 43._--Forelimbs erupted; larval mouthparts absent; corner of +mouth between nostril and eye; transverse bands present on hind limbs; +tail greatly reduced (about 8 mm. in length). + +_Stage 44._--Sacral hump barely noticeable; tail reduced to a stub; +corner of mouth at level of pupil of eye; dorsal surfaces pale +olive-green; venter white. + +Changes proceed in a definite pattern during the growth and development +of tadpoles. Larval teeth are absent in hatchlings; the inner tooth-rows +develop first, and the third lower row last. At metamorphosis the third +lower row is the first to be lost. The tail increases gradually in +length relative to the body. In stage 25 the tail is 52.1 per cent of +the total length, and in stage 36, 64.6 per cent. In later stages the +tail becomes relatively shorter through resorption. Duellman and Klaas +(1964:320) noted a great size-variation in _Triprion_ tadpoles in stage +25. No such variation is apparent in any stage of any of the species of +_Smilisca_ studied. + +The growth and development of the other species of _Smilisca_ do not +differ significantly from that of _S. phaeota_. The tadpoles of _S. +sila_ and _sordida_ from streams have relatively longer tails at +hatching. For example, in tadpoles of _S. sordida_ the average length of +tail is 64.0 per cent of the body-length in stage 25, and in stage 37, +67.0 per cent. + +_Behavior_ + +The tadpoles of _S. baudini_, _cyanosticta_, _phaeota_, and _puma_ are +pelagic inhabitants of shallow ponds. Early stages of _S. baudini_ in +which external gills are present have been observed to hang vertically +with the gills spread out at the surface of the water, a behavior noted +by Zweifel (1964:206) in tadpoles of _Phrynohyas venulosa_, which also +develop in warm, standing water having a relatively low oxygen-tension. +When disturbed the pelagic tadpoles usually dive and seek shelter amidst +vegetation or in mud on the bottom. This behavior was observed in _S. +baudini_, _cyanosticta_, and _phaeota_ by day and at night. No tadpoles +of _S. puma_ were observed by day; those seen at night were near the +surface of small water-filled depressions in a grassy marsh; they +responded to light by taking refuge in the dense grass. Perhaps tadpoles +of this species are negatively phototactic and remain hidden by day. + +The stream-inhabiting tadpoles of _S. sila_ and _sordida_ live in clear +pools in rocky streams, where they were observed to cling by their +mouths to rocks in the stream and to seek shelter amidst pebbles or +beneath rocks and leaves on the bottom. These tadpoles are not found in +shallow riffles. + +We have not found tadpoles of two species of _Smilisca_ in the same body +of water and therefore cannot offer observations on ecological +relationships in sympatric situations. + + + + +PHYLOGENETIC RELATIONSHIPS + + +Identifiable hylid remains are known from the Miocene to the Recent, but +these fossils are mostly fragmentary and provide little useful +information regarding the phylogenetic relationships of living genera. +Frogs of the genus _Smilisca_ are generalized and show no striking +adaptations, either in their structure or in their modes of life +history. + + +Interspecific Relationships + +In attempting to understand the relationships of the species of +_Smilisca_ we have emphasized osteological characters. The phylogeny +suggested by these characters is supported by other lines of evidence, +including external morphology, tadpoles, and breeding calls. + +Our concept of the prototype of the genus _Smilisca_ is a moderate-sized +hylid having: (1) a well-developed frontoparietal fontanelle, +(2) frontoparietal lacking lateral processes, (3) no bony +squamosal-maxillary arch, (4) a fully ossified ethmoid, (5) paired +subgular vocal sac, (6) moderately webbed fingers and toes, (7) +relatively few supernumerary tubercles on the digits, (8) eggs deposited +in clumps in ponds, (9) anteroventral mouth in tadpoles bordered by one +row of labial papillae, but median part of upper lip bare, (10) tail +relatively short and deep in tadpoles, and (11) a breeding call +consisting of a series of like notes. + +Two phyletic lines evolved from this prototype. The first of these was +the stock that gave rise to the _baudini_ group. The evolutionary +changes that took place in this line included increase in size, +development of a lateral curvature of the maxillary, and an increased +amount of cranial ossification, especially in the dermal roofing bones. +This phyletic line retained the larval characters and breeding call of +the prototype. The second phyletic line gave rise to the _sordida_ group +and diverged from the prototype in the development of an angular +maxillary and a breeding call consisting of a primary note followed by +secondary notes. The frogs in this phyletic line retained the moderate +size of the prototype and did not develop additional dermal bone. Our +concept of the phylogenetic relationships is shown graphically in Figure +17. + +Within the _baudini_ group one stock retained separate nasals and did +not develop a bony squamosal-maxillary arch, but broad lateral processes +developed on the frontoparietals. The tadpoles remained unchanged from +the primitive type. This stock evolved into _S. phaeota_. In the other +stock the nasals became fully ossified and a bony squamosal-maxillary +arch developed. One branch of this second stock retained tadpoles having +only one row of labial papillae and did not develop lateral processes on +the frontoparietals; this branch evolved into _S. cyanosticta_. The +other branch diverged and gave rise to _S. baudini_ by developing +relatively shorter hind legs, large lateral processes on the +frontoparietals, and tadpoles having two rows of labial papillae. + +Within the _sordida_ group the cranial features remained unchanged in +one line, which gave rise to _S. sila_, whereas in a second line the +nasals were reduced, and their long axes shifted with the result that +they are not parallel to the maxillaries; the amount of ossification of +the ethmoid was reduced, and the tadpoles developed two rows of labial +papillae. In this second line one branch retained the pond-breeding +habits and gave rise to _S. puma_, whereas a second branch became +adapted to stream-breeding and gave rise to _S. sordida_. + + _baudini_ + \ _cyanosticta_ + \ / + + _phaeota_ _sordida_ + \ / / + + puma_ / + \ \/ _sila_ + \ /_____/ + \ / + \ / + \ / + | + | + Prototype + + [Illustration: FIG. 17. Hypothesized phylogenetic relationships + of the species of _Smilisca_.] + +Certain aspects of this proposed phylogeny warrant further comment. +Features such as the deposition of additional bone that roofs the skull +or that forms lateral projections from the frontoparietals, like those +in _S. baudini_ and _phaeota_, are minor alterations of dermal elements +and not basic modifications of the architecture of the skull. +Consequently, we hypothesize the independent development of these dermal +changes in _S. baudini_ and _phaeota_. Similar kinds of dermal +modifications have evolved independently in many diverse groups of +frogs. + +Likewise, we propose the parallel development of stream-adapted tadpoles +in _S. sordida_ and _sila_; in both cases the tadpoles adapted to +changing environmental conditions (see following section on evolutionary +history). Tadpoles of _S. sordida_ already had two rows of labial +papillae before entering the streams; subsequently the tadpoles +developed complete rows of papillae, ventral mouths and long tails +having low fins. Possibly the tadpoles of _S. sila_ had two rows of +labial papillae prior to their adapting to stream conditions; in the +process of adapting they developed ventral mouths and long tails having +low fins. Similar modifications in tadpoles have occurred in many +diverse groups of Middle American hylids, such as _Plectrohyla_, +_Ptychohyla_, the _Hyla uranochroa_ group, and the _Hyla taeniopus_ +group. + +Our lack of concern about coloration is due to the fact that, with the +exception of the blue spots on the flanks and posterior surfaces of the +thighs in some species, the coloration of _Smilisca_, consisting of a +pattern of irregular dark marks on a paler dorsum and dark transverse +bars on the limbs, is not much different from that of many other +Neotropical hylids. Blue is a structural color, rare among Amphibia, +which is achieved by the absence of lipophores above the guanophores. +Thus, the incident light rays at the blue end of the spectrum are +reflected by the guanophores without interference by an overlying yellow +lipophore screen. According to Noble (1931), lipophores are capable of +amoeboid movement that permits shifts in their positions, between or +beneath the guanophores. We do not know whether this behavior of +lipophores is widespread and is effected in response to environmental +changes, or whether it is a genetically controlled attribute that is +restricted in appearance. If the latter is the case we must assume that +the prototype of _Smilisca_ possessed such an attribute which was lost +in _S. baudini_, _phaeota_, and _puma_. The development of blue spots is +not constant in _S. sordida_ and _S. sila_; in _S. cyanosticta_ the +spots range in color from blue to pale green. + +The coloration of the tadpoles is not distinctive, except for the +presence of dorsal blotches on the tails of _S. sila_ and _sordida_. +However, the similarity in pattern cannot be interpreted as indicating +close relationships because nearly identical patterns are present in +_Hyla legleri_ and some species of _Prostherapis_. This disruptive +coloration seems to be directly associated with the pebble-bottom, +stream-inhabiting tadpoles. + +In the _baudini_ group, _S. phaeota_ and _cyanosticta_ are allopatric, +whereas _S. baudini_ occurs sympatrically with both of those species. +The call of _S. baudini_ differs notably from the calls of _S. phaeota_ +and _cyanosticta_, which are more nearly alike. Although in the +phylogenetic scheme proposed here _S. sila_ is considered to be more +distantly related to _S. puma_ than is _S. sordida_, the calls of _S. +sila_ and _puma_ more closely resemble one another than either resembles +that of _S. sordida_. _Smilisca sila_ and _puma_ are allopatric, whereas +_S. sordida_ is broadly sympatric with both of those species. We assume +that in their respective phyletic lines the differentiation of both _S. +baudini_ and _sordida_ was the result of genetic changes in +geographically isolated populations. Subsequently, each species +dispersed into areas inhabited by other members of their respective +groups. Selection for differences in the breeding calls helped to +reinforce other differences in the populations and thereby aided in +maintaining specificity. + + +Evolutionary History + +With respect to temporal and spatial aspects of evolution in _Smilisca_, +we have tried to correlate the phylogenetic evidence on _Smilisca_ with +the geologic data on Middle America presented by Lloyd (1963), Vinson +and Brineman (1963), Guzman and Cserna (1963), Maldonado-Koerdell +(1964), and Whitmore and Stewart (1965). Likewise, we have borne in mind +the evidence for, and ideas about, the evolution of the Middle American +herpetofauna given by Dunn (1931b), Schmidt (1943), Stuart (1950, 1964) +Duellman (1958, MS), and Savage (MS). + +According to Stuart's (1950) historical arrangement of the herpetofauna, +_Smilisca_ is a member of the Autochthonous Middle American Faunal +Element, and according to Savage's (MS) arrangement the genus belongs to +the Middle American Element, a fauna which was derived from a +generalized tropical American unit that was isolated in tropical North +America by the inundation of the Isthmian Link in early Tertiary, that +developed _in situ_ in tropical North America, and that was restricted +to Middle America by climatic change in the late Cenozoic. + +Savage (MS) relied on the paleogeographic maps of Lloyd (1963) to +hypothesize the extent and centers of differentiation of the Middle +American Faunal Element. According to Lloyd's concept, Middle America in +the Miocene consisted of a broad peninsula extending southeastward to +about central Nicaragua, separated from the Panamanian Spur of +continental South America by shallow seas. A large island, the Talamanca +Range, and remnants of the Guanarivas Ridge formed an archipelago in the +shallow sea. The recent discovery of remains of mammals having definite +North American affinities in the Miocene of the Canal Zone (Whitmore and +Stewart, 1965) provides substantial evidence that at least a peninsula +was continuous southeastward from Nuclear Central America to the area of +the present Canal Zone in early mid-Miocene time. South America was +isolated from Central America by the Bolivar Trough until late +mid-Pliocene. + +Thus, in the mid-Tertiary the broad peninsula of Nuclear Central +America, which consisted of low and moderately uplifted regions having a +tropical mesic climate, provided the site for the evolution of +_Smilisca_. It is not possible to determine when the genus evolved, but +to explain the differentiation of the species it is unnecessary to have +the ancestral _Smilisca_ present prior to the Miocene. + +We view the Miocene _Smilisca_ as the prototype described in the +preceding section, and suppose that it lived in the mesic tropical +environment of the eastern part of the Central American Peninsula (in +what is now Costa Rica and western Panama). Two stocks differentiated, +probably in middle Miocene times; one of these, the ancestral stock of +the _baudini_ group, was widespread on the Caribbean lowlands from the +Nicaraguan Depression to the Bolivar Trough, and the other, the +ancestral stock of the _sordida_ group, was restricted to the Pacific +lowlands of the same region. In late Miocene time the ancestral stock of +the _baudini_ group dispersed northwestward around the deep embayment in +the Nicaraguan depression into upper Central America (in what is now +Honduras and Guatemala) and thence into southern Mexico. Apparently +differentiation took place on each side of the Nicaraguan Depression; +the frogs to the south of the depression evolved into _S. phaeota_, +whereas those to the north of the depression represented the stock from +which _S. baudini_ and _cyanosticta_ arose. Prior to the uplift of the +mountains in the late Miocene and the Pliocene the _baudini-cyanosticta_ +stock probably was widespread in northwestern Central America. The +elevation of the mountains resulted in notable climatic changes, +principally the development of sub-humid environments on the Pacific +lowlands. The frogs living on the Pacific lowlands became adapted to +sub-humid conditions and developed into _S. baudini_. The stock on the +Caribbean lowlands remained in mesic environments and evolved into _S. +cyanosticta_. + +Possibly in the middle Miocene before the Talamanca Range in Costa Rica +and western Panama was greatly uplifted, the ancestral stock of the +_sordida_ group invaded the Caribbean lowlands of what is now Costa +Rica. The subsequent elevation of the Talamanca Range in the Pliocene +effectively isolated the ancestral stock of _S. sila_ on the Pacific +lowlands from the _puma-sordida_ stock on the Caribbean lowlands. The +former was subjected to the sub-humid conditions which developed on the +Pacific lowlands when the Talamanca Range was uplifted. It adapted to +the sub-humid environment by living along streams and evolving +stream-adapted tadpoles. On the Caribbean side of the Talamanca Range +the _puma-sordida_ stock inhabited mesic environments. The stock that +evolved into _S. puma_ remained in the lowlands as a pond-breeding frog, +whereas those frogs living on the slopes of the newly elevated mountains +became adapted for their montane existence by developing stream-adapted +tadpoles and thus differentiated into _S. sordida_. + +Probably the six species of _Smilisca_ were in existence by the end of +the Pliocene; at that time a continuous land connection existed from +Central America to South America. The climatic fluctuations in the +Pleistocene, and the post-Wisconsin development of present climatic and +vegetational patterns in Middle America, brought about the present +patterns of distribution of the species. From its place of origin on the +Caribbean lowlands of lower Central America, _S. phaeota_ dispersed +northward into Nicaragua and southward along the Pacific slopes of +northwestern South America. Perhaps in the late Pleistocene or in +post-Wisconsin time when mesic conditions were more widespread than now, +_S. phaeota_ moved onto the Pacific lowlands of Costa Rica. Its route +could have been through the Arenal Depression. Subsequent aridity +restricted its range on the Pacific lowlands to the Golfo Dulce region. +Climatic fluctuation in northern Central America restricted the +distribution of _S. cyanosticta_ to mesic habitats on the slopes of the +Mexican and Guatemalan highlands and to certain humid areas on the +lowlands. _Smilisca baudini_ was well adapted to sub-humid conditions, +and the species dispersed northward to the Rio Grande Embayment and to +the edge of the Sonoran Desert and southward into Costa Rica. In +southern Mexico and Central America the species invaded mesic habitats. +Consequently, in some areas it is sympatric with _S. cyanosticta_ and +_phaeota_. + +_Smilisca puma_ dispersed northward onto the Caribbean lowlands of +southern Nicaragua. Its southward movements probably were limited by the +ridges of the Talamanca Range that extend to the Caribbean coast in the +area of Punta Cahuita in Costa Rica. _Smilisca sila_ dispersed along the +Pacific lowlands and slopes of the mountains from eastern Costa Rica and +western Panama through eastern Panama to northern Colombia. Climatic +fluctuation in the Pleistocene evidently provided sufficient altitudinal +shifts in environments in the Talamanca Range to permit _S. sordida_ to +move onto the Pacific slopes. From its upland distribution the species +followed streams down to both the Caribbean and Pacific lowlands, where +it is sympatric with _S. puma_ on the Caribbean lowlands and _S. sila_ +on the Pacific lowlands. + +The evolution of the species-groups of _Smilisca_ was effected through +isolation by physical barriers in the Cenozoic; the differentiation of +the species was initiated by further isolation of populations by changes +in physiography and climate. Present patterns of distribution resulted +from Pleistocene and post-Wisconsin climatic changes. Today, sympatric +species have different breeding habits and breeding calls which +reinforce the differences in morphology. + + + + +SUMMARY AND CONCLUSIONS + + +The genus _Smilisca_ is composed of six species of tree frogs; each +species is defined on the basis of adult morphology, larval characters, +and breeding behavior. Keys are provided to aid in the identification of +adults and of tadpoles. + +Analysis of the characters and examination of type specimens indicates +that several currently-recognized taxa are synonymous, as follows: + + 1. _Hyla beltrani_ Taylor, 1942 = _Smilisca baudini_. + 2. _Hyla gabbi_ Cope, 1876 = _Smilisca sordida_. + 3. _Hyla manisorum_ Taylor, 1954 = _Smilisca baudini_. + 4. _Hyla nigripes_ Cope, 1876 = _Smilisca sordida_. + 5. _Hyla wellmanorum_ Taylor, 1952 = _Smilisca puma_. + +_Smilisca phaeota cyanosticta_ Smith, 1953 is elevated to specific rank, +and one new species, _Smilisca sila_, is named and described. + +The skeletal system of developmental stages and the adult of _Smilisca +baudini_ is described, and the skull is compared with that of other +members of the genus. + +The tadpoles are described, compared, and illustrated; the larval +development of _Smilisca phaeota_ is described. + +Breeding behavior and breeding calls are described and compared. Some +species of _Smilisca_ have breeding choruses. Two species, _S. sila_ and +_sordida_, breed in streams, whereas the others breed in ponds. + +The genus is considered to be part of the Middle American Faunal +Element; the species are thought to have differentiated in response to +ecological diversity and historical opportunities provided by Cenozoic +changes in physiography and climate. + + + + +LITERATURE CITED + + +BAIRD, S. F. + + 1854. Descriptions of new genera and species of North American frogs. + Proc. Acad. Nat. Sci. Philadelphia, 7:59-62. April 27. + + 1859. Reptiles of the boundary. United States and Mexican boundary + survey. Washington, D. C., p. 35, pl. 41. + + +BALDAUF, R. J. + + 1959. Morphological criteria and their use in showing bufonid phylogeny. + Jour. Morph., 104:527-560. May. + + +BARBOUR, T. + + 1923. Notes on reptiles and amphibians from Panama. Occas. Papers + Mus. Zool. Univ. Michigan, 129:1-16. January 25. + + +BLAIR, W. F. + + 1959. Call structure and species groups in U. S. treefrogs (_Hyla_). + Southwest. Nat., 3:77-89. June 1, 1959. + + 1962. Non-morphological data in anuran classification. Syst. Zool., + 11:72-84. June. + + 1963. Evolutionary relationships of North American toads of the genus + Bufo: A progress report. Evolution, 17:1-16. March. + + +BOGERT, C. M. + + 1960. The influence of sound on the behavior of amphibians and reptiles. + _in_ Lanyon, W. E. and Tavolga, W. N. (Eds.) Animal sounds and + communication, pp. 137-320. + + +BRATTSTROM, B. H. + + 1957. The phylogeny of the Salientia based on skeletal morphology. Syst. + Zool., 6:70-74. June. + + +BREDER, C. M., Jr. + + 1946. Amphibians and reptiles of the Rio Chucunaque drainage, Darien, + Panama, with notes on their life histories and habits. Bull. + Amer. Mus. Nat. Hist., 86:375-436, pls. 42-60. August 26. + + +BROCCHI, P. + + 1877. Note sur quelques batrachiens hylaeformes recuilles au Mexique + et au Guatemala. Bull. Soc. Philom. Paris, ser. 7, 1:122-132. + + +CARVALHO, A. L. + + 1954. A preliminary synopsis of the genera of American microhylid frogs. + Occas. Papers Mus. Zool. Univ. Michigan, 555:1-19, pl. 1. July 16. + + +COCHRAN, D. M. + + 1961. Type specimens of reptiles and amphibians in the United States + National Museum. Bull. U. S. Nat. Mus., 220:xv + 291 pp. + + +COPE, E. D. + + 1862. Catalogues of the reptiles obtained during the explorations of + the Parana, Paraguay, Vermejo and Uraguay rivers.... Proc. + Acad. Nat. Sci. Philadelphia, 14, pt. 9:346-359. + + 1865. Third contribution to the herpetology of tropical America. Proc. + Acad. Nat. Sci. Philadelphia, 17:185-198. October. + + 1871. Ninth contribution to the herpetology of tropical America. Proc. + Acad. Nat. Sci. Philadelphia, 23, pt. 2:200-224. + + +COPLAND, S. J. + + 1957. Australian tree frogs of the genus _Hyla_. Proc. Linnean Soc. New + South Wales, 82, pt. 1:9-108. September. + + +DUELLMAN, W. E. + + 1956. The frogs of the hylid genus _Phrynohyas_ Fitzinger, 1843. Misc. + Publ. Mus. Zool. Univ. Michigan, 96:1-47, pls. 1-6. February 21. + + 1958. A monographic study of the colubrid snake genus _Leptodeira_. + Bull. Amer. Mus. Nat. Hist., 114:1-152, pls. 1-31. February 24. + + 1963a. A review of the Middle American tree frogs of the genus + Ptychohyla. Univ. Kansas Publ. Mus. Nat. Hist., 15:297-349, + pls. 11-18. October 18. + + 1963b. A new species of tree frog, genus _Phyllomedusa_, from Costa Rica. + Rev. Biol. Trop., 11(1):1-23. October. + + 1964. A review of the frogs of the Hyla bistincta group. Univ. Kansas + Publ. Mus. Nat. Hist., 15:469-491. March 2. + + 1965. Frogs of the _Hyla taeniopus_ group. Copeia, no. 2:159-168. + June 25. + + +DUELLMAN, W. E. and KLAAS, L. T. + + 1964. The biology of the hylid frog _Triprion petasatus_. Copeia, + no. 2:308-321. June 30. + + +DUMERIL, A. M. C. and BIBRON, G. + + 1841. Erpetologie Generale ou histoire naturelle complete des reptiles, + vol. 8, 792 pp. + + +DUNN, E. R. + + 1931a. The amphibians of Barro Colorado Island. Occas. Papers Boston + Soc. Nat. Hist., 5:403-421. October 10. + + 1931b. The herpetological fauna of the Americas. Copeia, no. 3:106-119. + October 30. + + 1944. Herpetology of the Bogota area. Rev. Acad. Colombiana Cien. + Exact., Fis. Nat., 6:68-81. + + +FOUQUETTE, M. J., JR. + + 1960. Isolating mechanisms in three sympatric tree frogs in the Canal + Zone. Evolution, 14:484-497. December. + + +FUNKHOUSER, ANNE + + 1957. A review of the neotropical tree-frogs of the genus + _Phyllomedusa_. Occas. Papers Nat. Hist. Mus. Stanford Univ., + 5:1-89. April 1. + + +GADOW, H. + + 1908. Through southern Mexico. London, Witherby and Co. xvi + 527 pp. + + +GAIGE, H. T., HARTWEG, N. and STUART, L. C. + + 1937. Notes on a collection of amphibians and reptiles from eastern + Nicaragua. Occas. Papers Mus. Zool. Univ. Michigan, 357:1-18. + October 26. + + +GOIN, C. J. + + 1961. Synopsis of the genera of hylid frogs. Ann. Carnegie Mus., + 36:5-18. July 14. + + +GOSNER, K. L. + + 1960. A simplified table for staging anuran embryos and larvae with + notes on identification. Herpetologica, 16:183-190. September 23. + + +GRIFFITHS, I. + + 1959. The phylogeny of _Sminthillus limbatus_ and the status of the + Brachycephalidae (Amphibia, Salientia). Proc. Zool. Soc. London, + 132:457-487, pls. 1-4. + + +GUZMAN, E. J. and CSERNA, Z. + + 1963. Tectonic History of Mexico. Amer. Assoc. Petrol. Geol., + Mem. 2:113-129. + + +HECHT, M. K. + + 1962. A reevaluation of the early history of the frogs. Part I. Syst. + Zool., 11:39-44. March. + + 1963. A reevaluation of the early history of the frogs. Part II. Syst. + Zool., 12:20-35. March. + + +JOHNSON, C. + + 1959. Genetic incompatibility in the call races of _Hyla versicolor_ + Le Conte in Texas. Copeia, no. 4:327-335. December 30. + + +LLOYD, J. J. + + 1963. Tectonic history of the south Central-American Orogen. Amer. + Assoc. Petrol. Geol., Mem. 2:88-100. + + +MALDONADO-KOERDELL, M. + + 1964. Geohistory and paleogeography of Middle America, _in_ Wauchope, + R. and West, R. C. (Eds.). Handbook of Middle American Indians, + vol. 1, Univ. Texas Press, Austin, 570 pp. + + +MASLIN, T. P. + + 1963. Notes on some anuran tadpoles from Yucatan, Mexico. Herpetologica, + 19:122-128. July 3. + + +MITTLEMAN, M. B. and LIST, J. C. + + 1963. The generic differentiation of the swamp treefrogs. Copeia, + no. 2:80-83. May 29. + + +NOBLE, G. K. + + 1931. The biology of the amphibia. McGraw Hill, New York, 577 pp. + + +ORTON, G. L. + + 1957. The bearing of larval evolution on some problems in frog + classification. Syst. Zool., 6:79-86. June. + + +PETERS, W. + + 1863. Mittheilungen uber neue Batrachier. Monats. Konigl. Akad. Wiss. + Berlin, pp. 445-471. + + 1873. Uber eine neue Schildkroetenart, _Cinosternon Effeldtii_ und einige + andere neue oder weniger bekannte Amphibien. Monats. Konigl. + Akad. Wiss. Berlin, pp. 603-618, pl. 5. October 16. + + +RIVERO, J. A. + + 1961. Salientia of Venezuela. Bull. Comp. Zool., 126:1-207. November. + + +SAVAGE, J. M. and CARVALHO, A. L. + + 1953. The family position of Neotropical frogs currently referred to the + genus _Pseudis_. Zoologica, 38:193-200. + + +SCHMIDT, K. P. + + 1941. The amphibians and reptiles of British Honduras. Zool. Ser. Field + Mus. Nat. Hist., 22:475-510. December 30. + + 1943. Corollary and commentary for "Climate and Evolution." Amer. + Midl. Nat., 30:241-253. July. + + +SCHMIDT, O. + + 1857. Diagnosen neuer Froesche des zoologischen Cabinets zu Krakau. + Sitzungb. Konigl. Akad. Wiss. Math.-Natur. Cl., 24(1):10-15. + March. + + 1858. Deliciae Herpetogicae Musei Zoologici Cracoviensis. Denkschr. + K. K. Akad. Wiss. Math.-Natur. Cl., 14(2):237-258, pls. 1-3. + + +SMITH, H. M. + + 1953. A new subspecies of the treefrog _Hyla phaeota_ Cope of Central + America. Herpetologica, 8:150-152. January 30. + + +SMITH, H. M. and TAYLOR, E. H. + + 1950. Type localities of Mexican reptiles and amphibians. Univ. Kansas + Sci. Bull., 33:313-380. March 20. + + +STARRETT, P. + + 1960. A redefinition of the genus _Smilisca_. Copeia, no. 4:300-304. + December 30. + + +STEBBINS, R. C. and HENDRICKSON, J. R. + + 1959. Field studies of amphibians in Colombia, South America. Univ. + California Publ. Zool., 56:497-540. February 17. + + +STOKELY, P. S. and LIST, J. C. + + 1954. The progress of ossification in the skull of the cricketfrog + _Pseudacris nigrita triseriata_. Copeia, no. 3:211-217. July 29. + + +STUART, L. C. + + 1935. A contribution to a knowledge of the herpetology of a portion of + the savanna region of central Peten, Guatemala. Misc. Publ. Mus. + Zool. Univ. Michigan, 29:1-56, pls. 1-4, 1 map. October 1. + + 1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. + Publ. Mus. Zool. Univ. Michigan, 69:1-109. June 12. + + 1950. A geographic study of the herpetofauna of Alta Verapaz, Guatemala. + Contr. Lab. Vert. Biol., 45:1-77, pls. 1-9, 1 map. May. + + 1954. Herpetofauna of the southeastern highlands of Guatemala. Contr. + Lab. Vert. Biol., 68:1-65, pls. 1-4. November. + + 1958. A study of the herpetofauna of the Uaxactun-Tikal area of northern + El Peten, Guatemala. Contr. Lab. Vert. Biol., 75:1-30. June. + + 1961. Some observations on the natural history of tadpoles of + _Rhinophrynus dorsalis_ Dumeril and Bibron. Herpetologica, + 17:73-79. July 11. + + 1964. Fauna of Middle America, _in_ Wauchope, R. and West, R. C. + (Eds.). Handbook of Middle American Indians, vol. 1, Univ. + Texas Press, Austin, 570 pp. + + +TAYLOR, E. H. + + 1942. New Caudata and Salientia from Mexico. Univ. Kansas Sci. Bull., + 28:295-323. November 15. + + 1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., + 35:577-942. July 1. + + 1954. Additions to the known herpetological fauna of Costa Rica with + comments on other species. No. I. Univ. Kansas Sci. Bull., + 36:597-639. June 1. + + +TAYLOR, E. H. and SMITH, H. M. + + 1945. Summary of the collections of amphibians made in Mexico under + the Walter Rathbone Bacon Traveling Scholarship. Proc. U. S. + Natl. Mus., 95:521-613, pls. 18-32. June 30. + + +TIHEN, J. A. + + 1962. Osteological observations on New World Bufo. Amer. Midl. Nat., + 67:157-183. January. + + 1965. Evolutionary trends in frogs. Amer. Zoologist, 5:309-318. + + +VINSON, G. L. and BRINEMAN, J. H. + + 1963. Nuclear Central America, hub of Antillean Transverse Belt. Amer. + Assoc. Petrol. Geol., Mem. 2:101-112. + + +WHITMORE, F. C., JR. and STEWART, R. H. + + 1965. Miocene mammals and Central American seaways. Science, + 148:180-185. April 9. + + +ZWEIFEL, R. G. + + 1956. Two pelobatid frogs from the Tertiary of North America and their + relationships to fossil and Recent forms. Amer. Mus. Novitates, + 1762:1-45. April 6. + + 1958. Results of the Archbold Expeditions. No. 78. Frogs of the Papuan + hylid genus _Nyctimystes_. Amer. Mus. Novitates, 1896:1-51. + July 22. + + 1964. Life history of _Phrynohyas venulosa_ (Salientia: Hylidae) in + Panama. Copeia, no. 1:201-208. March 26. + + +_Transmitted March 14, 1966._ + + + + + [] + 31-3430 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + +MUSEUM OF NATURAL HISTORY + + +Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in a +particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. When +individuals request copies from the Museum, 25 cents should be included, +for each 100 pages or part thereof, for the purpose of defraying the +costs of wrapping and mailing. For certain longer papers an additional +amount, indicated below, toward some of the costs of production, is to +be included. + + * An asterisk designates those numbers of which the Museum's supply + is exhausted. + + Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. + + *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. + Pp. 1-444, 140 figures in text. April 9, 1948. + + Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, + and distribution. By Rollin H. Baker. Pp. 1-359, + 16 figures in text. June 12, 1951. + + *2. A quantitative study of the nocturnal migration of + birds. By George H. Lowery, Jr. Pp. 361-472, 47 + figures in text. June 29, 1951. + + 3. Phylogeny of the waxwings and allied birds. By M. Dale + Arvey. Pp. 473-530, 49 figures in text, 13 tables. + October 10, 1951. + + *4. Birds from the state of Veracruz, Mexico. By George H. + Lowery, Jr. and Walter W. Dalquest. Pp. 531-649, + 7 figures in text, 2 tables. October 10, 1951. + + Index. Pp. 651-681. + + *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, + 41 plates, 31 figures in text. December 27, 1951. + + Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. + + *Vol. 6. (Complete) Mammals of Utah, _taxonomy and distribution_. By + Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. + August 10, 1952. + + Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. + + Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. + + Vol. 9. Nos. 1-23 and index. Pp. 1-690, 1955-1960. + + Vol. 10. Nos. 1-10 and index. Pp. 1-626, 1956-1960. + + Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. + + Vol. 12. *1. Functional morphology of three bats: Eumops, Myotis, + Macrotus. By Terry A. Vaughan. Pp. 1-153, pls. 1-4, + 24 figures in text. July 8, 1959. + + *2. The ancestry of modern Amphibia: a review of the + evidence. By Theodore H. Eaton, Jr. Pp. 155-180, + 10 figures in text. July 10, 1959. + + 3. The baculum in microtine rodents. By Sidney Anderson. + Pp. 181-216, 49 figures in text. February 19, 1960. + + *4. A new order of fishlike Amphibia from the Pennsylvanian + of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou + Stewart. Pp. 217-240, 12 figures in text. May 2, 1960. + + 5. Natural history of the Bell Vireo, Vireo bellii Audubon. + By Jon C. Barlow. Pp. 241-296, 6 figures in text. + March 7, 1962. + + 6. Two new pelycosaurs from the lower Permian of Oklahoma. + By Richard C. Fox. Pp. 297-307, 6 figures in text. + May 21, 1962. + + 7. Vertebrates from the barrier island of Tamaulipas, + Mexico. By Robert K. Selander, Richard F. Johnston, + B. J. Wilks, and Gerald G. Raun. Pp. 309-345, pls. 5-8. + June 18, 1962. + + 8. Teeth of edestid sharks. By Theodore H. Eaton, Jr. + Pp. 347-362, 10 figures in text. October 1, 1962. + + 9. Variation in the muscles and nerves of the leg in two + genera of grouse (Tympanuchus and Pedioecetes). + By E. Bruce Holmes. Pp. 363-474, 20 figures in text. + October 25, 1963. $1.00. + + 10. A new genus of Pennsylvanian fish (Crossopterygii, + Coelacanthiformes) from Kansas. By Joan Echols. + Pp. 475-501, 7 figures in text. October 25, 1963. + + 11. Observations on the Mississippi kite in southwestern + Kansas. By Henry S. Fitch. Pp. 503-519. October 25, 1963. + + 12. Jaw musculature of the Mourning and White-winged doves. + By Robert L. Merz. Pp. 521-551, 22 figures in text. + October 25, 1963. + + 13. Thoracic and coracoid arteries in two families of birds, + Columbidae and Hirundinidae. By Marion Anne Jenkinson. + Pp. 553-573, 7 figures in text. March 2, 1964. + + 14. The breeding birds of Kansas. By Richard F. Johnston. + Pp. 575-655, 10 figures in text. May 18, 1964. 75 cents. + + 15. The adductor muscles of the jaw in some primitive + reptiles. By Richard C. Fox. Pp. 657-680, 11 figures in + text. May 18, 1964. + + Index. Pp. 681-694. + + Vol. 13. 1. Five natural hybrid combinations in minnows (Cyprinidae). + By Frank B. Cross and W. L. Minckley. Pp. 1-18. + June 1, 1960. + + 2. A distributional study of the amphibians of the Isthmus + of Tehuantepec, Mexico. By William E. Duellman. + Pp. 19-72, pls. 1-8, 3 figures in text. August 16, 1960. + 50 cents. + + 3. A new subspecies of the slider turtle (Pseudemys scripta) + from Coahuila, Mexico. By John M. Legler. Pp. 73-84, + pls. 9-12, 3 figures in text. August 16, 1960. + + *4. Autecology of the copperhead. By Henry S. Fitch. + Pp. 85-288, pls. 13-20, 26 figures in text. + November 30, 1960. + + 5. Occurrence of the garter snake, Thamnophis sirtalis, in + the Great Plains and Rocky Mountains. By Henry S. Fitch + and T. Paul Maslin. Pp. 289-308. 4 figures in text. + February 10, 1961. + + 6. Fishes of the Wakarusa River in Kansas. By James E. + Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure in + text. February 10, 1961. + + 7. Geographic variation in the North American cyprinid fish, + Hybopsis gracilis. By Leonard J. Olund and Frank B. + Cross. Pp. 323-348, pls. 21-24, 2 figures in text. + February 10, 1961. + + 8. Descriptions of two species of frogs, genus Ptychohyla; + studies of American hylid frogs, V. By William E. + Duellman. Pp. 349-357, pl. 25, 2 figures in text. + April 27, 1961. + + 9. Fish populations, following a drought, in the Neosho and + Marais des Cygnes rivers of Kansas. By James Everett + Deacon. Pp. 359-427, pls. 26-30, 3 figures in text. + August 11, 1961. 75 cents. + + 10. North American recent soft-shelled turtles (Family + Trionychidae). By Robert G. Webb. Pp. 429-611, + pls. 31-54, 24 figures in text. February 16, 1962. $2.00. + + Index. Pp. 613-624. + + + Vol. 14. 1. Neotropical bats from western Mexico. By Sydney Anderson. + Pp. 1-8. October 24, 1960. + + 2. Geographical variation in the harvest mouse, + Reithrodontomys megalotis, on the central Great Plains + and in adjacent regions. By J. Knox Jones, Jr., and B. + Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961. + + 3. Mammals of Mesa Verde National Park, Colorado. By Sydney + Anderson. Pp. 29-67, pls. 1 and 2, 3 figures in text. + July 24, 1961. + + 4. A new subspecies of the black myotis (bat) from eastern + Mexico. By E. Raymond Hall and Ticul Alvarez. Pp. 69-72, + 1 figure in text. December 29, 1961. + + 5. North American yellow bats, "Dasypterus," and a list of + the named kinds of the genus Lasiurus Gray. By E. + Raymond Hall and J. Knox Jones, Jr. Pp. 73-98, 4 figures + in text. December 29, 1961. + + 6. Natural history of the brush mouse (Peromyscus boylii) + in Kansas with description of a new subspecies. + By Charles A. Long. Pp. 99-110, 1 figure in text. + December 29, 1961. + + 7. Taxonomic status of some mice of the Peromyscus boylii + group in eastern Mexico, with description of a new + subspecies. By Ticul Alvarez. Pp. 111-120, 1 figure in + text. December 29, 1961. + + 8. A new subspecies of ground squirrel (Spermophilus + spilosoma) from Tamaulipas, Mexico. By Ticul Alvarez. + Pp. 121-124. March 7, 1962. + + 9. Taxonomic status of the free-tailed bat, Tadarida + yucatanica Miller. By J. Knox Jones, Jr., and Ticul + Alvarez. Pp. 125-133, 1 figure in text. March 7, 1962. + + 10. A new doglike carnivore, genus Cynarctus, from the + Clarendonian, Pliocene, of Texas. By E. Raymond Hall and + Walter W. Dalquest. Pp. 135-138. 2 figures in text. + April 30, 1962. + + 11. A new subspecies of wood rat (Neotoma) from northeastern + Mexico. By Ticul Alvarez. Pp. 139-143. April 30, 1962. + + 12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox + Jones, Jr., Ticul Alvarez, and M. Raymond Lee. + Pp. 145-159, 1 figure in text. May 18, 1962. + + 13. A new bat (Myotis) from Mexico. By E. Raymond Hall. + Pp. 161-164, 1 figure in text. May 21, 1962. + + *14. The mammals of Veracruz. By E. Raymond Hall and Walter + W. Dalquest. Pp. 165-362, 2 figures in text. May 20, + 1963. $2.00. + + 15. The Recent mammals of Tamaulipas, Mexico. By Ticul + Alvarez. Pp. 363-473, 5 figures in text. May 20, 1963. + $1.00. + + 16. A new subspecies of the fruit-eating bat, Sturnira + ludovici, from western Mexico. By J. Knox Jones, Jr., + and Gary L. Phillips. Pp. 475-481, 1 figure in text. + March 2, 1964. + + 17. Records of the fossil mammal Sinclairella, Family + Apatemyidae, from the Chadronian and Orellan. By William + A. Clemens, Jr. Pp. 483-491, 2 figures in text. + March 2, 1964. + + 18. The mammals of Wyoming. By Charles A. Long. Pp. 493-758, + 82 figures in text. July 6, 1965. $3.00. + + Index. Pp. 759-784. + + + Vol. 15. 1. The amphibians and reptiles of Michoacan, Mexico. By + William E. Duellman. Pp. 1-148, pls. 1-6, 11 figures in + text. December 20, 1961. $1.50. + + 2. Some reptiles and amphibians from Korea. By Robert G. + Webb, J. Knox Jones, Jr., and George W. Byers. + Pp. 149-173. January 31, 1962. + + 3. A new species of frog (genus Tomodactylus) from western + Mexico. By Robert G. Webb. Pp. 175-181, 1 figure in text. + March 7, 1962. + + 4. Type specimens of amphibians and reptiles in the Museum + of Natural History, The University of Kansas. By William + E. Duellman and Barbara Berg. Pp. 183-204. + October 26, 1962. + + 5. Amphibians and reptiles of the rainforests of southern + El Peten, Guatemala. By William E. Duellman. Pp. 205-249, + pls. 7-10, 6 figures in text. October 4, 1963. + + 6. A revision of snakes of the genus Conophis (Family + Colubridae, from Middle America). By John Wellman. + Pp. 251-295, 9 figures in text. October 4, 1963. + + 7. A review of the Middle American tree frogs of the genus + Ptychohyla. By William E. Duellman. Pp. 297-349, + pls. 11-18, 7 figures in text. October 18, 1963. + 50 cents. + + 8. Natural history of the racer Coluber constrictor. + By Henry S. Fitch. Pp. 351-468, pls. 19-22, 20 figures + in text. December 30, 1963. $1.00. + + 9. A review of the frogs of the Hyla bistincta group. + By William E. Duellman. Pp. 469-491, 4 figures in text. + March 2, 1964. + + 10. An ecological study of the garter snake, Thamnophis + sirtalis. By Henry S. Fitch. Pp. 493-564, pls. 23-25, + 14 figures in text. May 17, 1965. + + 11. Breeding cycle in the ground skink, Lygosoma laterale. + By Henry S. Fitch and Harry W. Greene. Pp. 565-575, + 3 figures in text. May 17, 1965. + + 12. Amphibians and reptiles from the Yucatan Peninsula, + Mexico. By William E. Duellman. Pp. 577-614, 1 figure + in text. June 22, 1965. + + 13. A new species of turtle, genus Kinosternon, from Central + America. By John M. Legler. Pp. 615-625, pls. 26-28, + 2 figures in text. June 20, 1965. + + 14. A biogeographic account of the herpetofauna of Michoacan, + Mexico. By William E. Duellman. Pp. 627-709, pls. 29-36, + 5 figures in text. December 30, 1965. + + 15. Amphibians and reptiles of Mesa Verde National Park, + Colorado. By Charles L. Douglas. Pp. 711-744, pls. 37 + and 38, 6 figures in text. March 7, 1966. + + Index in preparation. + + + Vol. 16. 1. Distribution and taxonomy of mammals of Nebraska. By J. + Knox Jones, Jr. Pp. 1-356, plates 1-4, 82 figures in + text. October 1, 1964. $3.50. + + 2. Synopsis of the lagomorphs and rodents of Korea. By J. + Knox Jones, Jr., and David H. Johnson. Pp. 357-407. + February 12, 1965. + + 3. Mammals from Isla Cozumel, Mexico, with description of + a new species of harvest mouse. By J. Knox Jones, Jr. + and Timothy E. Lawlor. Pp. 409-419, 1 figure in text. + April 13, 1965. + + 4. The Yucatan deer mouse, Peromyscus yucatanicus. + By Timothy E. Lawlor. Pp. 421-438, 2 figures in text. + July 20, 1965. + + 5. Bats from Gautemala. By J. Knox Jones, Jr. Pp. 439-472. + April 18, 1966. + + More numbers will appear in volume 16. + + + Vol. 17. 1. Localities of fossil vertebrates obtained from the + Niobrara Formation (Cretaceous) of Kansas. By David + Bardack. Pp. 1-14. January 22, 1965. + + 2. Chorda tympani branch of the facial nerve in the middle + ear of tetrapods. By Richard C. Fox. Pp. 15-21. + June 22, 1965. + + 3. Fishes of the Kansas River System in relation to + zoogeography of the Great Plains. By Artie L. Metcalf. + Pp. 23-189, 4 figures in text, 51 maps. March 24, 1966. + + 4. Factors affecting growth and production of channel + catfish, Ictalurus punctatus. By Bill A. Simco and + Frank B. Cross. Pp. 191-256, 13 figures in text. + June 6, 1966. + + 5. A new species of fringe-limbed tree frog, genus Hyla, + from Darien, Panama. By William E. Duellman. Pp. 257-262, + 1 figure in text. June 17, 1966. + + 6. Taxonomic notes on some Mexican and Central American + hylid frogs. By William E. Duellman. Pp. 263-279. + June 17, 1966. + + 7. Neotropical hylid frogs, genus Smilisca. By William E. + Duellman and Linda Trueb. Pp. 281-375, pls. 1-12, + 17 figures in text. July 14, 1966. + + More numbers will appear in volume 17. + + + + +Transcriber's Note + +With the exception of the corrections listed below and several minor +corrections not listed, the text presented is that which appeared in the +original printed version. The list of Kansas University publications has +been compiled at the end of the article. + + +Typographical Corrections + + Page Correction + ==== ================== + 287 cleared => cleaned + 292 Data based of => Data based on + 298 CNMH => CNHM + 299 Acahuitzotla => Acahuizotla + 304 cyanostica => cyanosticta + 305 Quatemala => Guatemala + 307 cyanostica => cyanosticta + 313 Matagalapa => Matagalpa + 322 Carribean => Caribbean + 323 Centralia => Centrali + 336 proportionaely => proportionately + 346 noticably => noticeably + 362 Fouquett => Fouquette + 372 CARVALJO => CARVALHO + 375 Dumeril => Dumeril + ii trutles => turtles + + + + + +End of the Project Gutenberg EBook of Neotropical Hylid Frogs, Genus Smilisca, by +William E. 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