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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

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

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.