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.

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 ( to 76 mm., to 90 mm.) having broad flat heads and a dark brown or black postorbital mark encompassing tympanum 2

Smaller frogs ( to 45 mm., 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