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UNIVERSITY OF KANSAS PUBLICATIONS

MUSEUM OF NATURAL HISTORY

Volume 18, No. 6, pp. 505-545, 7 figs., 4 pls.

December 2, 1969

The Systematics of the Frogs of the Hyla rubra Group in Middle America

JUAN R. LEÓN

University of Kansas Lawrence 1969

UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editors: Frank B. Cross, Philip S. Humphrey, Robert M. Mengel.

Volume 18, No. 6, pp. 505-545, 7 figs., 4 pls. Published December 2, 1969

UNIVERSITY OF KANSAS Lawrence, Kansas

PRINTED BY ROBERT R. (BOB) SANDERS, STATE PRINTER TOPEKA, KANSAS 1969

The Systematics of the Frogs of the Hyla rubra Group in Middle America

JUAN R. LEÓN

CONTENTS

PAGE

INTRODUCTION 508 Acknowledgments 508 Materials and Methods 509

THE HYLA RUBRA GROUP 509 Key to Species and Subspecies 510 Key to Known Tadpoles 511

ACCOUNTS OF SPECIES AND SUBSPECIES 511 Hyla boulengeri (Cope) 511 Hyla foliamorta Fouquette 520 Hyla rubra Laurenti 524 Hyla elaeochroa Cope 525 Hyla staufferi Cope 532 Hyla staufferi staufferi Cope 537 Hyla staufferi altae Dunn 540

EVOLUTIONARY HISTORY 540

LITERATURE CITED 543

INTRODUCTION

The tree frogs of the Hyla rubra group are abundant and form a conspicuous element of the Neotropical frog fauna. Representatives of the group occur from lowland México to Argentina; the greatest diversity is reached in the lowlands of southeastern Brazil (Cochran, 1955). The group apparently originated in South America; the endemic Central American species evolved from stocks that invaded Middle America after the closure of the Colombian Portal in the late Pliocene.

Dunn (1933) partially defined the rubra group as it occurs in Central America. Cope (1865, 1876, 1887), Brocchi (1881), Boulenger (1882), Günther (1901), Noble (1918), Kellogg (1932), Dunn and Emlen (1932), Stuart (1935), and Gaige (1936) dealt with the Middle American species now considered to make up the rubra group. More recently, Taylor (1952, 1958), Fouquette (1958), Starrett (1960), and Duellman (1960, 1963, 1966a) studied aspects of the taxonomy and biology of the species of this group. The five species of the rubra group in Central America have received ten different names. One species, Hyla staufferi, has received five names (two subspecies are recognized herein). Hyla boulengeri was named in the genus Scytopis, but the type species of Scytopis is a member of the genus Phrynohyas Fitzinger, 1843 (Duellman, 1956.)

Little has been published concerning the ecology, life history, osteology, and mating calls of the Middle American species of this group. The purpose of the present report is to describe the species occurring in Middle America and to comment on their distributions, ecology, cranial osteology, and mating calls, and in so doing provide evidence for the evolutionary history of the species inhabiting Middle America.

Acknowledgments

For permission to examine specimens in their care, I am grateful to Drs. Richard G. Zweifel, American Museum of Natural History (AMNH); Robert F. Inger, Field Museum of Natural History (FMNH); Ernest E. Williams, Museum of Comparative Zoology (MCZ); Hobart M. Smith, University of Illinois Museum of Natural History (UIMNH); Charles F. Walker, University of Michigan Museum of Zoology (UMMZ); Jay M. Savage, University of Southern California (USC); James A. Peters, United States National Museum (USNM); Richard J. Baldauf, Texas Cooperative Wildlife Collection (TCWC); and W. Frank Blair, Texas Natural History Collection (TNHC). KU refers to specimens in the Museum of Natural History, University of Kansas. For the loan of tape-recordings of mating calls I thank Drs. W. Frank Blair, University of Texas, and Richard G. Zweifel, American Museum of Natural History.