_Description and Variation._--Considerable variation in size

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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.

Neotropical Hylid Frogs, Genus Smilisca · 第 1 / 12 段