Chapter I: Part 1

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

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.

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.]

A

B

Skull of adult female _Smilisca baudini_ (KU 68184): (A) Dorsal;
(B) Ventral. x4.5.]

A

B

C

Skull of adult female _Smilisca baudini_ (KU 68184): (A) Lateral;
(B) Dorsal view of left mandible; (C) Posterior. x4.5.]

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.]

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.]

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.]

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.]

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.]

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.]

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).]

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).]

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.

_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.