Neotropical Hylid Frogs, Genus Smilisca

Part 1

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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

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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.

Part 2

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.

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