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Title: Neotropical Hylid Frogs, Genus Smilisca
Author: Duellman, William E., 1930-, Trueb, Linda
Language: English
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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_ (Duméril 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 México 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, Günther 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 José, 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 México, 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 Panamá under the
auspices of a grant from the National Institutes of Health (NIH
GM-12020) in cooperation with the Gorgas Memorial Laboratory in Panamá.


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 México 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_
     Duméril 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_ Duméril 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 México and Central
America, in some places to elevations of nearly 2000 meters, southward
from southern Sonora and Río Grande Embayment of Texas, including such
continental islands as Isla Cozumel, México, and Isla Popa and Isla
Cebaco, Panamá, to northern South America, where known from Caribbean
coastal regions and valleys of Río Cauca and Río 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= (Duméril and Bibron)

  _Hyla baudini_ Duméril and Bibron, Erpétologie général, 8:564, 1841
     [Holotype.--MNHN 4798 from "Mexico;" Baudin collector]. Günther,
     Catalogue Batrachia Salientia in British Museum, p. 105, 1858.
     Brocchi, Mission scientifique au Mexique ..., pt. 3, sec. 2, Études
     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. Günther, 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 México, 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_]. Diáz de León, Indice de los batracios
     que se encuentran en la República Mexicana, p. 20, June 1904.

  _Hyla muricolor_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
     14(9):359, 1862 [Holotype.--USNM 25097 from Mirador, Veracruz,
     México; 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 Panzós, Alta Verapaz, Guatemala; M.
     Bocourt collector]; Mission scientifique au Mexique ..., pt. 3,
     sec. 2, Études 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, México; A. Magaña 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 Batán, Limón 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 México, including Tamaulipas, Veracruz, the Yucatán
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)
                              |            |              |
  Córdoba, 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)
                              |            |              |
  Chichén-Itzá, Yucatán       | 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)
                              |            |              |
  Chinajá, 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)
                              |            |              |
  Limón, 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 México the average ratio varies from
45.6 in Veracruz to 47.6 on Isla del Carmen, Campeche. Specimens from
the Yucatán 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 Limón, 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 Chinajá,
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 México,
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 Matatán, 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
Cobán 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 Limón Province, Costa Rica (KU 34927 from
Batán 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 Petén
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 México 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
Petén, 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 Petén,
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 Chinajá, 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, México, _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 Chinajá,
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, Yucatán (KU 71720), and
4 kilometers west-southwest Puerto Juárez, Quintana Roo, México (KU
71721), whereas others, notably from 17 kilometers northeast of
Juchatengo, Oaxaca, México (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 Pisté, Yucatán, México. These
specimens are heavily pigmented like specimens that we have examined
from the Yucatán 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 Petén, 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, México, 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. Duméril and Bibron (1841:564) proposed the name
_Hyla baudini_ for a specimen (MNHN 4798) from México. Smith and Taylor
(1950:347) restricted the type locality to Córdoba, Veracruz, México, 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, México, 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. ×23;
    (B) young (KU 85438), snout-vent length 32.1 mm. ×9.]

    [Illustration: PLATE 2

    A

    B

    Skull of adult female _Smilisca baudini_ (KU 68184): (A) Dorsal;
    (B) Ventral. ×4.5.]

    [Illustration: PLATE 3

    A

    B

    C

    Skull of adult female _Smilisca baudini_ (KU 68184): (A) Lateral;
    (B) Dorsal view of left mandible; (C) Posterior. ×4.5.]

    [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). ×3.]

    [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). ×3.]

    [Illustration: PLATE 6

    A

    B

    C

    Living _Smilisca_: (A) _S. baudini_ (UMMZ 115179) from 1.7 km. W
    Xicotencatl, Tamaulipas, México; (B) _S. cyanosticta_ (UMMZ 118163)
    from Volcán San Martín, Veracruz, México; (C) _S. phaeota_
   (KU 64282) from Barranca del Río Sarapiquí, 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 Volcán, Chiriquí, Panamá;
    (C) _S. sordida_ (KU 64257) from 20 km. WSW San Isidro el General,
    San José 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_, Río 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). ×3.2.]

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 Batán,
Limón, 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 Río
Esnápe, Darién, Panamá.

    [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
México from southern Sonora and the Río 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 Solistahuacán, Chiapas, elevation 1675 meters,
and 10 kilometers northwest of Comitán, 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.

MEXICO: =Campeche=: Balchacaj, CNHM 102285, 102288, 102291, 102311,
UIMNH 30709-22, 30726; Champotón, UMMZ 73172 (2), 73176, 73180; 16 km. E
Champotón, UMMZ 73181; 5 km. S Champotón, KU 71369-75; 9 km. S
Champotón, KU 71367-8; 10.5 km. S Champotón, KU 71365-6, 71722
(tadpoles), 71723 (yg.); 24 km. S Champotón, UMMZ 73177 (2); Chuina, KU
75101-3; Ciudad del Carmen, UIMNH 30703-8; Dzibalchén, KU 75413-31;
Encarnación, CNHM 102282, 102289, 102294-5, 102300, 102306-8, 102312,
102314, 102316-7, 102319, 102322, UIMNH 30727-40, 30836-7; 1 km. W
Escárcega, KU 71391-6; 6 km. W Escárcega, KU 71397-403; 7.5 km. W
Escárcega, KU 71376-89; 14 km. W Escárcega, KU 71390; 13 km. W, 1 km. N
Escárcega, KU 71404; 3 km. N Hopelchén, KU 75410-11; 2 km. NE Hopelchén,
KU 75412; Matamoras, CNHM 36573; Pitál, UIMNH 30741; 1 km. SW Puerto
Real, Isla del Carmen, KU 71345-64; San José Carpizo, UMMZ 99879; Tres
Brazos, CNHM 102284, UIMNH 30723-5; Tuxpeña Camp, UMMZ 73239.

=Chiapas=: Acacoyagua, USNM 114487-92; 2 km. W Acacoyagua, USNM
114493-4; 5 km. E Arroyo Minas, UIMNH 9533-7; Berriozabal, UMMZ 119186
(7); Chiapa de Corzo, UMMZ 119185 (2); Cintalapa, UIMNH 50077; Colonia
Soconusco, USNM 114495-9; 5 km. W Colonia Soconusco, UMMZ 87885 (7);
Comitán, UMMZ 94438; 10 km. NW Comitán, KU 57185; El Suspiro, UMMZ
118819 (11); Escuintla, UMMZ 88271 (7), 88278, 88327, 109233; 6 km. NE
Escuintla, UMMZ 87856 (26); 3 km. E Finca Juárez, UIMNH 9538; Finca
Prussia, UMMZ 95167; Honduras, UMMZ 94434-7; La Grada, UMMZ 87862;
21 km. S La Trinitaria, UIMNH 9540-1; 14.4 km. SW Las Cruces, KU
64239-44; Palenque, UIMNH 49286, USNM 114473-84; 2 km. NW Pueblo Nuevo
Solistahuacán, KU 57182-4, UMMZ 119948 (8), 121514; 1.3 km. N Puerto
Madero, KU 57186-9; 4 km. N Puerto Madero, KU 57190-1; 8 km. N Puerto
Madero, UMMZ 118379 (2); 12 km. N Puerto Madero, KU 57192; 17.6 km. N
Puerto Madero, UMMZ 118378; Rancho Monserrata, UIMNH 9531-2, UMMZ
102266-7; Region Soconusco, UIMNH 33542-56; San Bartola, UIMNH 9519-30;
San Gerónimo, UIMNH 30804; San Juanito, USNM 114485-6; San Ricardo, CNHM
102406; Solosuchiapa, KU 75432-3; Tapachula, CNHM 102208, 102219,
102239, 102405, UIMNH 25046, 30802-3; Tonolá, AMNH 531, CNHM 102232,
102416, UIMNH 30805-9, USNM 46760; Tuina, KU 41593 (skeleton); Tuxtla
Gutierrez, CNHM 102231, 102248; 6 km. E Tuxtla Gutierrez, UIMNH 9539;
10 km. E Tuxtla Gutierrez, UMMZ 119949.

=Chihuahua=: 2.4 km. SW Toquina, KU 47226-7; Riito, KU 47228.

=Coahuila=: mountain near Saltillo, UIMNH 30833-4.

=Colima=: No specific locality, CNHM 1632; Colima, AMNH 510-11;
Hacienda Albarradito, UMMZ 80029 (2); Hacienda del Colomo, AMNH 6208;
Los Mezcales, UMMZ 80028; Manzanillo, AMNH 6207, 6209; Paso del Río,
CNHM 102207, 102229-30, UIMNH 30819-21, UMMZ 110875 (3); Periquillo,
UMMZ 80025 (3), 80026 (14); 1.6 km. SW Pueblo Juárez, UMMZ 115564;
Queseria, CNHM 102204, 102216-7, 102224, UIMNH 30816-8, UMMZ 80023 (7),
80024 (7); Santiago, UMMZ 80027; 7.2 km. SW Tecolapa, UMMZ 115184.

=Guerrero=: Acahuizotla, UF 1338 (2), 1339-40, UMMZ 119182 (2), 119184;
3 km. S Acahuizotla, KU 87183-7; Acapulco, AMNH 55276, UMMZ 121879 (4),
USNM 47909; 3 km. N Acapulco, UMMZ 110127; 8 km. NW Acapulco, UF 11203
(7); 27 km. NE Acapulco, UIMNH 26597-610; Agua del Obispo, CNHM 102214,
102290, 102293, 102310, 102413, KU 60413, 87180-2, UIMNH 30764-6;
Atoyca, KU 87175-8; Buena Vista, CNHM 102279, 102304, 102313, 102315,
UIMNH 30774; Caculutla, KU 87179; 20 km. S Chilpancingo, CNHM 102242,
102401, 102410-1, 102415; Colonia Buenas Aires, UMMZ 119189; El
Limoncito, CNHM 102292, 102303, 102321, 102414; El Treinte, CNHM 102212,
102221, 102237, 102240-1, UIMNH 30783-5, USNM 114508-10; Laguna Coyuca,
UMMZ 80960 (2); 3 km. N Mazatlán, UIMNH 30777-9; 9 km. S Mazatlán, CNHM
102209, 102215, 102234, 102246, UIMNH 30781-2; Mexcala, CNHM 102399,
102403, 102409, 106539-40, UIMNH 30775-6; Ocotito, KU 60414-23; 5.4 km.
N Ocotito, UMMZ 119181 (4); 1.6 km. N Organos, UIMNH 30752-63; Palo
Blanco, CNHM 102283, 102286, 102305, 102320, 102404, UIMNH 30767-70; Pie
de la Cuesta, AMNH 55275, 59202-5; Puerto Marquéz, AMNH 59200-1 (13);
5.6 km. S San Andreas de la Cruz, KU 87173-4; San Vincente, KU 87172;
Zaculapán, UMMZ 119183.

=Hidalgo=: Below Tianguistengo, CNHM 102318.

=Jalisco=: Atenqueque, KU 91435-6; 5 km. NE Autlán, UIMNH 30810; 5 km. E
Barro de Navidad, UMMZ 110900; Charco Hondo, UMMZ 95247; Puerto
Vallarta, UIMNH 41346; between La Huerta and Tecomates, KU 91437; 3 km.
SE La Resolana, KU 27619, 27620 (skeleton); 11 km. S, 1.6 km. E
Yahualica, KU 29039; Zapotilitic, CNHM 102238.

=Michoacán=: Aguililla, UMMZ 119179 (5); Apatzingán, CNHM 38766-90, KU
69101 (skeleton); 7 km. E Apatzingán, UMMZ 112843; 11 km. E Apatzingán,
UMMZ 112841 (3); 27 km. S Apatzingán, KU 37621-3; 1.6 km. N Arteaga,
UMMZ 119180; Charapendo, UMMZ 112840; Coahuayana, UMMZ 104458; El
Sabino, CNHM 102205-6, 102210-1, 102220, 102228, 102233, UIMNH 30822-3;
La Placita, UMMZ 104456; La Playa, UMMZ 105163; 30 km. E Nueva Italia,
UMMZ 120255 (2); 4 km. S Nueva Italia, UMMZ 112842; Ostula, UMMZ 104457
(4); Salitre de Estopilas, UMMZ 104459; San José de la Montaña, UMMZ
104461 (2); 11 km. S Tumbiscatio, KU 37626; 12 km. S Tzitzio, UMMZ
119178.

=Morelos=: 3.5 km. W Cuautlixco, KU 87188-90; 1 km. NE Puente de Ixtla,
KU 60393-4; 20 km. S Puente de Ixtla, CNHM 102400, UIMNH 30832;
Tequesquitengo, AMNH 52036-9.

=Nayarit=: 3 km. S Acaponeta, UMMZ 123030 (4); 56 km. S Esquinapa
(Sinaloa), KU 73909; Jesús María, AMNH 58239; San Blas, KU 28087, 37624,
62360-2, USNM 51408; 8.6 km. E San Blas, UMMZ 115185; Tepic, UIMNH
30812-5; 4 km. E Tuxpan, KU 67786; 11 km. SE Tuxpan, UIMNH 7329-31,
7335-59.

=Nuevo León=: Galeana, CNHM 34389; Salto de Cola de Caballo, CNHM
30628-31, 30632 (40), 30633-7, 34454-67.

=Oaxaca=: 11 km. S Candelaria, UIMNH 9515-8; Cerro San Pedro, 24 km. SW
Tehuantepec, UMMZ 82156; Chachalapa, KU 38199; 8 km. S Chiltepec, KU
87191; 12 km. S Chivela, UMMZ 115182; Coyul, USNM 114512; Garza Mora,
UIMNH 40967-8; Juchatengo, KU 87193; 17 km. NE Juchatengo, KU 87645
(tadpoles), 87646 (young); Juchitán, USNM 70400; Lagartero, UIMNH 9514;
Matías Romero, AMNH 52143-5; 25 km. N Matías Romero, KU 33822-8; 7 km. S
Matías Romero, UIMNH 42703; Mirador, AMNH 6277, 13832-9, 13842-55; Mira
León, 1.6 km. N Huatulco, UIMNH 9503-4; Mixtequillo, AMNH 13924;
Pochutla, KU 57167-81, UIMNH 9505-13; Quiengola, AMNH 51817, 52146; Río
del Corte, UIMNH 48677; Río Mono Blanco, UIMNH 36831; Río Sarabia, 5 km.
N Sarabia, UMMZ 115180 (4); 2.5 km. N Salina Cruz, KU 57165-6; San
Antonio, UIMNH 37286; 5 km. NNW San Gabriel Mixtepec, KU 87192; San
Pedro del Istmo, UIMNH 37197; Santo Domingo, USNM 47120-2; 3.7 km. N
Sarabia, UMMZ 115181 (3); Tapanatepec, KU 37793 (skeleton), 37794, UIMNH
9542, UMMZ 115183; between Tapanatepec and Zanatepec, UIMNH 42704-25;
Tecuane, UMMZ 82163 (3); Tehuantepec, AMNH 52625, 52639, 53470, UMMZ
82157-8, 82159 (9), 82160 (4), 82161 (8), 82162 (12), 112844-5, 118703,
USNM 10016, 30171-4, 30188; 4.5 km. W Tehuantepec, KU 59801-12
(skeletons), 69102-3 (skeletons); 10 km. S Tehuantepec, KU 57163-4;
Temazcal, USC 8243 (3); 3 km. S Tolocita, KU 39666-9; Tolosa, AMNH
53605; Tuxtepec, UMMZ 122098 (2); 2 km. S Valle Nacional, KU 87194-5;
11 km. N Vista Hermosa, KU 87196, 87639-41 (tadpoles), 87642-3 (young),
87644 (tadpoles); Yetla, KU 87197.

=Puebla=: 16 km. SW Mecatepec (Veracruz), UIMNH 3657-8; San Diego, AMNH
57714, USNM 114511; Vegas de Suchil, AMNH 57712; Villa Juárez, UF 11205.

=Quintana Roo=: Cóba, CNHM 26937; Esmeralda, UMMZ 113551; 4 km. NNE
Felipe Carrillo Puerto, KU 71417-8; Pueblo Nuevo X-Can, KU 71405; 10 km.
ENE Pueblo Nuevo X-Can, KU 71406; 4 km. WSW Puerto Juárez, KU 71407-11,
71721 (tadpoles); 12 km. W Puerto Juárez, KU 71412-6; San Miguel, Isla
de Cozumel, UMMZ 78542 (6), 78543 (10), 78544 (2); 3.5 km. N San Miguel,
Isla de Cozumel, KU 71419-22; 10 km. E San Miguel, Isla de Cozumel, UMMZ
78541; Telantunich, CNHM 26950.

=San Luis Potosí=: Ciudad Valles, AMNH 57776-81 (12), CNHM 37193,
102297, KU 23705; 21 km. N Ciudad Valles, UMMZ 118377; 6 km. E Ciudad
Valles, UF 3524; 24 km. E Ciudad Valles, UF 7340 (2); 5 km. S Ciudad
Valles, UIMNH 30751; 16 km. S Ciudad Valles, AMNH 52953; 30 km. S Ciudad
Valles, CNHM 102394, 102402, 102412, UIMNH 30749-50; 63 km. S Ciudad
Valles, UIMNH 19247-58; Pujal, UMMZ 99872 (2); Río Axtla, near Axtla,
AMNH 53211-5, 59516, KU 23706; Tamazunchale, AMNH 52675, CNHM 39621-2,
102226, 102281, UF 7615 (2), UIMNH 26596, UMMZ 99506 (9), 118701 (2),
USNM 114468; 17 km. N Tamazunchale, UIMNH 3659; 2.4 km. S Tamazunchale,
AMNH 57743; 17 km. E Tamuin, UF 11202 (2); Xilitla, UIMNH 19259-60.

=Sinaloa=: 8 km. N. Carrizalejo, KU 78133; 4 km. NE Concordia, KU 73914;
5 km. SW Concordia, KU 75438-9; 6 km. E Cosalá, KU 73910; Costa Rica,
16 km. S. Culiacán, UIMNH 34887-9; 51 km. SSE Culiacán, KU 37792; El
Dorado, KU 60392; 1.6 km. NE El Fuerte, CNHM 71468; Isla Palmito del
Verde, middle, KU 73916-7; 21 km. NNE Los Mochis, UIMNH 40536-7;
Matatán, KU 73913; 7.3 km. SW Matatán, KU 78464, 78466-70; Mazatlán,
AMNH 12562, UMMZ 115197 (3); 57 km. N Mazatlán, UIMNH 38364; Plomosas,
USNM 47439-40; Presidio, UIMNH 30811, USNM 14082; Rosario, KU 73911-2;
5 km. E Rosario, UIMNH 7360-76; 8 km. SSE Rosario, KU 37625; 5 km. SW San
Ignacio, KU 78465; 1.6 km. ENE San Lorenzo, KU 47917-24; Teacapán, Isla
Palmito del Verde, KU 73915; 9.6 km. NNW Teacapán, KU 91410; Villa
Unión, KU 78471; 9 km. NE Villa Unión, KU 75434-7; 1 km. W Villa Unión,
AMNH 59284.

=Sonora=: Guiracoba, AMNH 51225-38 (25).

=Tabasco=: 4 km. NE Comalcalco, AMNH 60313; Teapa, UMMZ 119943; 5 km. N
Teapa, UMMZ 119940, 119944, 122997 (2); 10 km. N Teapa, UMMZ 119187,
119188 (2); 13 km. N Teapa, UMMZ 119941 (2), 119945 (3), 120254 (2);
21 km. N Teapa, UMMZ 119942, 119947; 29 km. N Teapa, UMMZ 119946 (11);
Tenosique, USNM 114505-7.

=Tamaulipas=: Acuña, UMMZ 99864; 5 km. S Acuña, UMMZ 101180; 13 km. N
Antiguo Morelos, UIMNH 40532-5; 3 km. S Antiguo Morelos, UF 11204; 3 km.
NE Chamal, UMMZ 102867; 1.6 km. E Chamal, UMMZ 110734; Ciudad Mante,
UMMZ 80957, 80958 (3), 106400 (3); 16 km. N Ciudad Victoria, CNHM
102408; 34 km. NE Ciudad Victoria, KU 60395-411; 8.8 km. S Ciudad
Victoria, UIMNH 19261-3; 11 km. W Ciudad Victoria, UIMNH 30924; 16 km. W
Ciudad Victoria, UIMNH 30825; 3 km. W El Carizo, UMMZ 111279; Gómez
Farías, UMMZ 110837-8; 8 km. NE Gómez Farías, UMMZ 102265, 102916 (4),
102917, 104110 (5), 105493, 110836 (2), 111274-7; 8 km. NW Gómez Farías,
UMMZ 101178 (7), 101179 (3), 101362-3, 101364 (2), 108799 (2), 110129,
111278, 111280; 8 km. W Gómez Farías, UMMZ 102859 (2); 16 km. W
Gonzales, KU 37795-6; Jiménez, KU 60412; La Clementina, 6 km. W Forlan,
USNM 106244; Limón, UIMNH 30831; Llera, USNM 140137-40; 3 km. E Llera,
UIMNH 16858; 21 km. S Llera, UIMNH 30828-9; 23 km. S Llera, UIMNH 30830;
11 km. SW Ocampo, UMMZ 118956; 22 km. W, 5 km. S Piedra, KU 37568-71;
Rio Sabinas, UMMZ 97976; 5 km. W San Gerardo, UMMZ 110733 (2); Santa
Barbara, UMMZ 111272-3; Villagrán, CNHM 102280, 102287, 102299, 102309,
UIMNH 30826-7; 1.7 km. W Xicotencatl, UMMZ 115179.

=Veracruz=: 1.6 km. NW Acayucan, UMMZ 115189; 28.5 km. SE Alvarado, UMMZ
119933; 2.4 km. SSW Amatitlán, UMMZ 115195; Barranca Metlac, UIMNH
38365; Boca del Río, UIMNH 26619-30, UMMZ 74954 (9); 16 km. S Boca del
Río, UIMNH 26631; between Boca del Río and Anton Lizardo, UIMNH 42701;
Canadá, CNHM 102397; Catemaco, UMMZ 118702 (4); Ciudad Alemán, UMMZ
119608 (3); Córdoba, CNHM 38665-7, USNM 30410-3; 5.2 km. ESE Córdoba, KU
71423-35, 89924 (skeleton); 7 km. ESE Córdoba, UMMZ 115176 (4);
Cosamaloapan, UMMZ 115193 (2); Coyame, UIMNH 36853-6, 38366, UMMZ 111461
(3), 111462-3; 1 km. SE Coyame, UMMZ 121202 (3); Cuatotolapam, UMMZ
41625-39; Cuautlapan, CNHM 38664, 70591-600, 102218, 102398, KU 26300,
26302, 26309, 26312-3, 26315-6, 26321, 26336, 26339, 26347 (skeleton),
26354, 55614-21 (skeletons), UIMNH 11236-67, 11269-71, 11273, 26611-8,
30792-5, UMMZ 85466 (6), 115173 (25), 115175 (7), USNM 114433-57; Dos
Ríos, CNHM 39623; 5 km. ENE El Jobo, KU 23843, 23845, 23847; 6.2 km. E
Encero, UIMNH 30835; Escamilo, CNHM 102298, UIMNH 30788; 1 km. N Fortín,
UF 11201; 4 km. SW Huatusco, UMMZ 115177; 1 km. SW Huatusco, UMMZ
123119; 10 km. SE Hueyapan, UMMZ 115190; 20 km. S Jesús Carranza, KU
23844, 23846, 27399; 38 km. SE Jesús Carranza, KU 23417; Laguna
Catemaco, UMMZ 119932 (62); 1.6 km. N La Laja, UIMNH 3651; La Oaxaqueña,
AMNH 43930-1; 17 km. E Martínez de la Torre, UIMNH 36630-2; 6.2 km. W
Martínez de la Torre, UIMNH 3652-4; Minatitlán, AMNH 52141-2; Mirador,
USNM 25097-8, 115178; 6 km. S Monte Blanco, UF 11200 (4); 21 km. E
Nanchital, UMMZ 123004; 2 km. S Naranja, UMMZ 115188 (3); 1.6 km. NE
Novillero, UMMZ 115194 (2); 3 km. NE Novillero, UMMZ 115196; 5.2 km. NE
Novillero, UMMZ 115192 (4); 6 km. NE Novillero, UMMZ 115191; 5 km. N
Nuevo Colonia, UMMZ 105066; Orizaba, USNM 16563-6; 4 km. NE Orizaba,
UMMZ 120251 (2); Panuco, UMMZ 118922; Paraje Nuevo, UMMZ 85465 (2),
85467 (35), 85468 (36); Paso del Macho, UIMNH 49281; Paso de Talaya,
Jicaltepec, USNM 32365, 84420; Pérez, CNHM 1686 (5); 20 km. N Piedras
Negras, Río Blanco, KU 23708; Plan del Río, KU 26310, 26333-5, 26345,
26354, UMMZ 102069, 102070 (5); Potrero, UIMNH 49282-5, UMMZ 88799,
88805, 88806 (2), USNM 32391-5; Potrero Viejo, CNHM 102296, KU 26301,
26304-5, 26307-8, 26311, 26317-20, 26323-25, 26326-8 (skeletons),
26329-31, 26332 (skeleton), 26337-8, 26340-4, 26346, 26348, 26351,
26353, 27400-12, UIMNH 30800, UMMZ 88800 (2), 88802 (15), 88803 (9),
88804, USNM 114458-67; 5 km. S Potrero Viejo, KU 26303, 26314, 26322;
Puente Nacional, UIMNH 21783-8; 3 km. N Rinconada, UMMZ 122099 (5); Río
de las Playas, USNM 118635-6; Río Seco, UMMZ 88801 (9); Rodriguez Clara,
CNHM 102225; San Andrés Tuxtla, CNHM 102213, 102222, 102227, 102247,
UIMNH 30789-91; 10 km. NW San Andrés Tuxtla, UMMZ 119935; 13.4 km. NW
San Andrés Tuxtla, UMMZ 119939 (2); 19.8 km. NW San Andrés Tuxtla, UMMZ
119938; 27.2 km. NW San Andrés Tuxtla, UMMZ 119936 (6); 48 km. NW San
Andrés Tuxtla, UMMZ 119937; 4 km. W San Andrés Tuxtla, UMMZ 115187; 37.4
km. S San Andrés Tuxtla, UMMZ 119934 (12); 15 km. ESE San Juan de la
Punta, KU 23707; San Lorenzo, USNM 123508-12; 3 km. SW San Marcias KU
23841; 1.5 km. S Santa Rosa, UIMNH 42702; 2 km. S Santiago Tuxtla, UMMZ
121201 (4); Sauzel, UMMZ 121239; 14 km. E Suchil, UIMNH 46880; 15 km. S
Tampico (Tamaulipas), UMMZ 103322 (4); 4 km. N Tapalapan, UMMZ 115186
(2); Tecolutla, UIMNH 42677-700; 16 km. NW Tehuatlán, UIMNH 3660-3;
5 km. S Tehuatlán, KU 23842; Teocelo, KU 26306; Tierra Colorado, CNHM
102393, 102395-6, UIMNH 30789-91; Veracruz, AMNH 6301-4, 59398-402,
UIMNH 30801, UMMZ 115174, 122060 (2); 24 km. W Veracruz, CNHM 104570-2.

=Yucatán=: No specific locality, CNHM 548, 49067, USNM 32298;
Chichén-Itzá, CNHM 20636, 26938-49, 36559-62, UIMNH 30742-6, UMMZ 73173
(6), 73174 (14), 73175 (14), 73178-9, 76171, 83107 (2), 83108, 83109
(2), 83915 (30), USNM 72744; 9 km. E Chichén-Itzá, KU 71438-9; 12 km. E
Chichén-Itzá, KU 71440; Mérida, CNHM 40659-66, UIMNH 30747-8, UMMZ
73182; 6 km. S Mérida, KU 75194; 8.8 km. SE Ticul, UMMZ 114296;
Valladolid, CNHM 26934-6; Xcalah-op, CNHM 53906-14; 3.5 km. E Yokdzonot,
KU 71441-3, 71720 (tadpoles).

BRITISH HONDURAS: Belize, CNHM 4153, 4384-5, 4387, UMMZ 75310, USNM
26065; Bokowina, CNHM 49064-5; Cocquercote, UMMZ 75331 (2); Cohune
Ridge, UMMZ 80738 (15); Double Falls, CNHM 49066; El Cayo, UMMZ 75311;
6 km. S El Cayo, MCZ 37856; Gallon Jug, MCZ 37848-55; Manatee, CNHM
4264-7; Mountain Pine Ridge, MCZ 37857-8; San Augustin, UMMZ 80739; San
Pedro, Columbia, MCZ 37860-2; Valentin, UMMZ 80735 (4), 80736 (2), 80737
(2); 5 km. S Waha Loaf Creek, MCZ 37859.

GUATEMALA: =Alta Verapaz=: 5.1 km. NE Campur, KU 68464 (tadpoles), 67465
(young); 28.3 km. NE Campur, KU 64203-22, 68183-4 (skeletons); Chamá,
MCZ 15792-3, UMMZ 90895 (7), 90896 (5), 90897 (29), 90898 (12), 90899;
Chinajá, KU 55939-41, 57193-8, 60018-20 (tadpoles), 60021 (eggs), 60022
(tadpoles); Cobán, CNHM 21006; Cubilquitz, UMMZ 90902 (10); Finca
Canihor, UMMZ 90908; Finca Chicoyou, KU 57246-8, 60026 (young), 64202,
68466-7 (tadpoles); Finca Los Alpes, KU 64197-201, 68463 (tadpoles);
Finca Los Pinales, UMMZ 90903 (2); Finca Tinajas, BYU 16031; Finca
Volcán, UMMZ 90905 (4), 90906-7; Panzos, MNHN 6313, UMMZ 90904; Samac,
UMMZ 90900; Samanzana, UMMZ 90901 (6).

=Baja Verapaz=: Chejel, UMMZ 90909 (7), 90910 (3); San Gerónimo, UMMZ
84076 (16).

=Chiquimula=: 1.6 km. SE Chiquimula, UMMZ 98112; Esquipulas, UMMZ
106793 (28).

=El Petén=: 20 km. NNW Chinajá (Alta Verapaz), KU 57199-240; Flores,
UMMZ 117985; La Libertad, KU 60024 (young), UMMZ 75313-20, 75323 (2),
75324 (7), 75325 (13), 75326 (2), 75327 (11), 75328 (12), 75329 (2);
3 km. SE La Libertad, KU 57243-4; 13 km. S La Libertad, MCZ 21458 (2);
Pacomon, USNM 71334; Piedras Negras, USNM 114469-71; Poptún, UMMZ
120475; Poza de la Jicotea, USNM 114672; Ramate-Yaxha trail, UMMZ 75321;
Río de la Pasión between Sayaxché and Subín, KU 57151; Río San Román,
16 km. NNW Chinajá (Alta Verapaz), KU 55942-6; Sacluc, USNM 25131;
Sayaxché, KU 57144-5; Tikal, UMMZ 117983 (7), 117984 (5), 117993 (5),
120474 (5); Toocog, KU 57241-2, 60023 (young), 60025 (young); Uaxactún,
UMMZ 70401-3; Yaxha, UMMZ 75322; 19 km. E Yaxha, UMMZ 75330 (4).

=El Quiché=: Finca Tesoro, UMMZ 89166 (3), 90549 (tadpoles).

=Escuintla=: Río Guacalate, Masagua, USNM 125239; Tiquisate,
UMMZ 98262 (7).

=Guatemala=: 16 km. NE Guatemala, KU 43545-53.

=Huehuetenango=: Finca San Rafael, 16 km. SE Barillas, CNHM 40912-6;
45 km. WNW Huehuetenango, KU 64223-4; Jacaltenango, UMMZ 120080 (6),
120081 (14), 120082 (13).

=Izabál=: 2 km. SW Puerto Matías de Gálvez, KU 60027-8 (tadpoles);
Quiriguá, CNHM 20587, UMMZ 70060.

=Jalapa=: Jalapa, UMMZ 98109, 106792 (11).

=Jutiapa=: Finca La Trinidad, UMMZ 107728 (10); Jutiapa, UMMZ 106789;
1.6 km. SE Mongoy, KU 43069; Santa Catarina Mita, UMMZ 106790.

=Progreso=: Finca Los Leones, UMMZ 106791.

=Quetzaltenango=: Coatepeque, AMNH 62204.

=Retalhueleu=: Casa Blanca, UMMZ 107725 (18); Champerico, UMMZ 107726
(3).

=San Marcos=: Talisman Bridge, USNM 128056-7.

=Santa Rosa=: Finca La Guardiana, UMMZ 107727 (6); Finca La Gloria, UMMZ
107724 (6); 1.6 km. WSW El Molino, KU 43065-8.

EL SALVADOR: =La Libertad=: 16 km. NW Santa Tecla, KU 43542-4.
=Morazán=: Divisadero, USNM 73284. =San Salvador=: San Salvador, CNHM
65087-99, KU 61955-88, 62138-9 (skeletons), 62154 (eggs), 62155-60
(tadpoles), 68462 (tadpoles), UMMZ 117586 (3), 118380 (3), USNM 140278.

HONDURAS: State unknown: Guaimas, UMMZ 58391. =Atlantidad=: Isla de
Roatán, CNHM 34551-4; La Ceiba, USNM 64985, 117589-91; Lancetilla, MCZ
16207-11; Tela, MCZ 15774-5, 28080, UMMZ 58418, USNM 82173-4.
=Choluteca=: 1.5 km. NW Choluteca, KU 64228-32; 10 km. NW Choluteca, KU
64233; 10 km. E Choluteca, KU 64226-7; 12 km. E Choluteca, KU 64225; 5
km. S Choluteca, USC 2700 (2). =Colón=: Bambú, UF 320; Belfate, AMNH
45692-5; Patuca, USNM 20261. =Comayagua=: La Misión, 3.5 leagues N
Siguatepeque, MCZ 26424-5. =Copán=: Copán, UMMZ 83026 (2). =Cortés=:
Cofradía, AMNH 45345-6; Hacienda Santa Ana, CNHM 4724-31; Lago de Yojoa,
MCZ 26410-1; Río Lindo, AMNH 54972. =El Paraiso=: El Volcán, MCZ 26436.
=Francisco Morazán=: Tegucigalapa, BYU 18301-4, 18837-41, MCZ 26395-7,
USNM 60499. =Gracias A Dios=: Río Segovia, MCZ 24543. =Santa Barbara=:
Santa Barbara, USNM 128062-5.

NICARAGUA: =Chinandega=: 4 km. N, 2 km. W Chichigalpa, KU 85385;
Chinandega, MCZ 2632; Río Tama, USNM 40022; San Antonio, KU 84944-9
(skeletons), 85386-403. =Chontales=: 1 km. NE Acoyapa, KU 64234.
=Estelí=: Finca Daraili, 5 km. N, 15 km. E Condega, KU 85404-8; Finca
Venecia, 7 km. N, 16 km. E Condega, KU 85409. =León=: 1.6 km. ENE
Poneloya, KU 43084-5. =Managua=: Managua, USNM 79989-90; 8 km. NW
Managua, KU 43094-110; 20 km. NE Managua, KU 85412; 21 km. NE Managua,
KU 85413-4; 5 km. SW Managua, KU 43086-93; 2 km. N Sabana Grande, KU
85411; 3 km. N Sabana Grande, KU 43070-8; 20 km. S, 0.5 km. W Tipitapa,
KU 85410. =Matagalpa=: Guasqualie, UMMZ 116493; Matagalpa, UMMZ 116492;
19 km. N Matagalpa, UMMZ 116494. =Río San Juan=: Greytown, USNM 19585-6,
19767-8. =Rivas=: Javillo, UMMZ 123001; Moyogalpa, Isla Ometepe, KU
85428-37, 87706 (tadpoles); Peñas Blancas, KU 85417; Río Javillo, 3 km.
N, 4 km. W Sapoá, KU 85418-20, 85438 (skeleton); 13.1 km. SE Rivas, KU
85415; 14.8 km. SE Rivas, KU 85421-3; 11 km. S, 3 km. E Rivas, KU 85416;
16 km. S Rivas, MCZ 29009-10; 7.7 km. NE San Juan del Sur, KU 85426-7;
16.5 km. NE San Juan del Sur, KU 85424-5, 87705 (young); 5 km. SE San
Pablo, KU 43079-83. =Zelaya=: Cooley, AMNM 7063-8, 8019-20, 8022,
8034-5; Cukra, AMNH 8016-7; Musahuas, Río Huaspuc, AMNH 58428-31; 11 km.
NW Rama, Río Siquia, UMMZ 79708, 79709 (5), 79710 (2); Río Escondido,
USNM 19766, 20701; Río Siquia at Río Mico, UMMZ 79707 (10); Sioux
Plantation, AMNH 7058-61, 8023-33.

COSTA RICA: =Alajuela=: Los Chiles, AMNH 54639; Orotina, MCZ 7960-1; San
Carlos, USNM 29991. =Guanacaste=: La Cruz, USC 8232 (3); 4.3 km. NE La
Cruz, UMMZ 123002; 18.4 km. S La Cruz, USC 8136; 23.5 km. S La Cruz, USC
8094 (4); 3 km. W La Cruz, USC 8233 (4); 2 km. NE Las Cañas, KU 64235-7;
Las Huecas, UMMZ 71212-3; Liberia, KU 36787, USC 8161; 11.5 km. N
Liberia, USC 8149; 13 km. N Liberia, USC 8139; 22.4 km. N Liberia, USC
8126; 8 km. NNW Liberia, KU 64238; 8.6 km. ESE Playa del Coco, USC 8137;
21.8 km. ESE Playa del Coco, USC 8138; Río Piedra, 1.6 km. W Bagaces,
USC 7027; Río Bebedero, 5 km. S Bebedero, KU 64158; 5 km. NE Tilarán, KU
36782-6. =Heredia=: 13 km. SW Puerto Viejo, KU 64142-6. =Limón=: Batán,
KU 34927; Guacimo, USC 621; Pandora, USC 505 (3); Suretka, KU 36788-9;
Tortugero, UF 7697, 10540-2. =Puntarenas=: Barranca, CNHM 35254-6; 15
km. WNW Barranca, KU 64155-7, UMMZ 118381; 18 km. WNW Barranca, UMMZ
118382 (4); 4 km. WNW Esparta, KU 64159-96, 68178-82 (skeletons); 19 km.
NW Esparta, KU 64147-54.


=Smilisca cyanosticta= (Smith), new combination

  _Hyla phaeota_, Taylor, Univ. Kansas Sci. Bull., 28(5):80, May 15,
     1942. Taylor and Smith, Proc. U. S. Natl. Mus., 95(3185):589, June
     30, 1945.

  _Hyla phaeota cyanosticta_ Smith, Herpetologica, 8:150, Jan. 30,
     1953 [Holotype.--USNM 111147 from Piedras Negras, El Petén,
     Guatemala; Hobart M. Smith collector]. Shannon and Werler, Trans.
     Kansas Acad. Sci., 58:386, Sept. 24, 1955. Poglayen and Smith,
     Herpetologica, 14:11, April 25, 1958. Cochran, Bull. U. S. Natl.
     Mus., 220:57, 1961. Smith, Illinois Biol. Mono., 32:25, May, 1964.

  _Smilisca phaeota cyanosticta_, Stuart, Misc. Publ. Mus. Zool.
     Univ. Michigan, 122:42, April 2, 1963. Duellman, Univ. Kansas Publ.
     Mus. Nat. Hist., 15(5):229, Oct. 4, 1963.

_Diagnosis._--Size moderately large ([M] 56.0 mm., [F] 70.0 mm.); skull
as long as wide; frontoparietal fontanelle large; narrow supraorbital
flanges having irregular margins anteriorly; large squamosal not in
contact with maxillary; tarsal fold moderately wide, full length of
tarsus; inner metatarsal tubercle moderately large, low, flat,
elliptical; hind limbs relatively long; tibia usually more than 52 per
cent of snout-vent length; labial stripe silvery-white; lips lacking
vertical bars; loreal region pale green; pale bronze-colored stripe from
nostril along edge of eyelid to point above tympanum narrow, bordered
below by narrow dark brown stripe from nostril to eye, and broad dark
brown postorbital mark encompassing tympanum and terminating above
insertion of arm; flanks, dark brown with large pale blue spots;
anterior and posterior surfaces of thighs dark brown with small
pale blue spots on thighs. (Foregoing combination of characters
distinguishing _S. cyanosticta_ from any other species in genus.)

_Description and Variation._--The largest males are from Piedras Negras,
El Petén, Guatemala, and they average 52.5 mm. in snout-vent length
whereas males from Los Tuxtlas, Veracruz, average 50.6 mm. and those
from northern Oaxaca 50.3 mm. The smallest breeding male has a
snout-vent length of 44.6 mm. The average ratio of tibia length to
snout-vent length is 54.8 per cent in males from Piedras Negras,
and 56.4 and 56.3 per cent in males from Los Tuxtlas and Oaxaca,
respectively. The only other character showing noticeable geographic
variation is the size of the tympanum. The average ratio of the diameter
of the tympanum to the diameter of the eye is 76.3 per cent in males
from Piedras Negras, 71.8 from Oaxaca, and 66.9 from Los Tuxtlas.

The dorsal ground color of _Smilisca cyanosticta_ is pale green to tan
and the venter is creamy white. The dorsum is variously marked with dark
olive-green or dark brown spots or blotches (Pl. 6B). An interorbital
dark bar usually is present. The most extensive dark area is a V-shaped
mark in the occipital region with the anterior branches not reaching the
eyelids; this mark is continuous, by means of a narrow mid-dorsal mark,
with an inverted V-shaped mark in the sacral region. In many specimens
this dorsal marking is interrupted, resulting in irregular spots. In
some specimens the dorsum is nearly uniform pale green or tan with a few
small, dark spots. The hind limbs are marked by dark transverse bands,
usually three or four each on the thigh and shank, and two or three on
the tarsus. The webbing on the feet is brown. The loreal region is pale
green, bordered above by a narrow, dark brown canthal stripe extending
from the nostril to the orbit, which is bordered above by a narrow,
bronze-colored stripe extending from the nostril along the edge of the
eyelid to a point above the tympanum. The upper lip is white. A broad
dark brown mark extends posteriorly from the orbit and encompasses the
tympanum to a point above the insertion of the forelimb. The flanks are
dark brown with many pale blue, rounded spots, giving the impression of
a pale blue ground color with dark brown mottling enclosing spots. The
anterior and posterior surfaces of the thighs are dark brown with many
small pale blue spots. The inner surfaces of the shank and tarsus are
colored like the posterior surfaces of the thighs. Pale blue spots are
usually present on the proximal segments of the second and third toes. A
distinct white stripe is present on the outer edge of the tarsus and
fifth toe; on the tarsus the white stripe is bordered below by dark
brown. A white stripe also is present on the outer edge of the forearm
and fourth finger. The anal region is dark brown, bordered above by a
narrow transverse white stripe. The throat in breeding males is dark,
grayish brown with white flecks.

No geographic variation in the dorsal coloration is evident. Specimens
from the eastern part of the range (Piedras Negras and Chinajá,
Guatemala) have bold, dark reticulations on the flanks enclosing large
pale blue or pale green spots, which fade to tan in preservative.
Specimens from Oaxaca and Veracruz characteristically have finer dark
reticulations on the flanks enclosing smaller blue spots; in many of
these specimens the ventrolateral spots are smallest and are white.

All living adults are easily recognized by the presence of pale, usually
blue, spots on the flanks and thighs. Individuals under cover by day
have a tan dorsum with dark brown markings. A hiding individual at
Chinajá, Alta Verapaz, Guatemala (KU 55936), had a pale tan dorsum when
found; later the dorsal color changed to chocolate brown. A pale green
patch was present below the eye; the spots on the posterior surfaces of
the thighs were pale blue, and those on the flanks were yellowish green.
A calling male obtained 10 kilometers north-northwest of Chinajá (KU
55934) was reddish brown when found at night; later the dorsal color
changed to pale tan. A green patch below the eye was persistent. Usually
these frogs are green at night. The coloration of an adult male (KU
87201) from 11 kilometers north of Vista Hermosa, Oaxaca, México, was
typical: "When calling dorsum pale green; later changed to dull
olive-green. Flanks dark brown with pale blue spots in axilla and groin
and bluish white flecks on mid-flank. Anterior and posterior surfaces of
thighs, inner surfaces of shanks, and median dorsal surfaces of tarsi
dark brown with blue spots. Canthal and postorbital stripes dark
chocolate brown; labial stripe creamy white. Forearm, tarsal, and anal
stripes pale cream-color. Throat dark brown with yellow flecks; belly
and ventral surfaces of limbs creamy buff; webs pinkish tan; iris deep
bronze, brown below pupil." (Duellman, field notes, June 24, 1964.)

Some individuals have both green and brown coloration in life. An
individual obtained at night on the south slope of Volcán San Martín,
Veracruz, México, had a pale tan dorsum changing peripherally to pale
green. The dorsal markings were dark brown and dark olive-green.

In contrast to the color-changes noted above, the labial region below
the eye is always pale green, and pale spots are always present on the
flanks and thighs in adults. The iris is invariably golden or bronze
above and darker, usually brown, below. Minute black flecks are present
on the iris, and in some individuals these flecks are so numerous that
the eye appears gray.

Recently metamorphosed young have pale tan flanks, and the posterior
surfaces of the thighs are orange-yellow; pale spots are absent. A
juvenile (KU 55935) from Chinajá, Alta Verapaz, Guatemala, having a
snout-vent length of 35.0 mm. was pale yellowish tan above with
olive-green markings; the flanks were dark brown with pale blue spots,
and the anterior and posterior surfaces of the thighs were uniform
bright tomato red. A juvenile (UMMZ 121298), 18.6 mm. in snout-vent
length, from the southeast slope of Volcán San Martín, Veracruz, México,
had pale tan flanks lacking blue spots, but had red thighs. Apparently
the ontogenetic changes in coloration proceed as follows: (1) flanks
pale tan and thighs orange-yellow, both lacking spots, (2) flanks pale
tan and thighs red, both lacking spots, (3) flanks dark brown with blue
spots and thighs red, lacking spots, and (4) flanks and thighs dark
brown, both having pale blue spots.

_Natural History._--_Smilisca cyanosticta_ inhabits humid tropical
forest and cloud forest from the lowlands to elevations of about 1200
meters in Los Tuxtlas and to about 900 meters in northern Oaxaca. In
these moist environments the frogs apparently are active throughout the
year. Active individuals have been obtained in January, July, and August
in Los Tuxtlas, Veracruz, in June and July in northern Oaxaca, in
February and March at Chinajá, Guatemala, and Taylor and Smith reported
(1945:589) activity in May at Piedras Negras, Guatemala. Calling males
were observed as follows; in a rain barrel 11 kilometers north of Vista
Hermosa, Oaxaca, México, on June 23-28, 1964; in a quiet pool in a
stream 8 kilometers south of Yetla, Oaxaca, México, in July, 1963 (Dale
L. Hoyt, personal communication); in and near springs flowing into a
stream at Dos Amates, Veracruz, México, on August 4, 1959 (Douglas
Robinson, personal communication); and in a water-filled depression in a
log 10 kilometers west-northwest of Chinajá, Guatemala, on March 13,
1960. Taylor and Smith (1945:589) reported that individuals were found
at night on the ground at the edge of temporary pools at Piedras Negras,
Guatemala, on May 28-29, 1939. A clasping pair was found on a rock at
the edge of a small stream on the south slope of Volcán San Martín,
Veracruz, México, on July 11, 1959 (Douglas Robinson, personal
communication).

Only one individual has been observed in a tree at night. In the
daytime, individuals were found in elephant ear plants (_Xanthosoma_) at
Chinajá, Guatemala.

The breeding call consists of one or two moderately short notes that are
lower pitched than those of _S. baudini_, but higher pitched than those
of _S. phaeota_. Each note has a duration of 0.25 to 0.45 seconds and is
repeated at intervals of one-half minute to several minutes. Each note
is a vibrant "waunk," having 110 to 180 pulses per second and dominant
frequency of 1600 to 2100 cycles per second (Pl. 10B).

Apparently the eggs are deposited as loose clumps in the water. In eggs
in the yolk plug stage of development, the diameter of the embryo is
about 2.3 mm.; that of the outer envelope is 4.0 mm. Hatchling tadpoles
have total lengths of 5.8 to 6.5 mm. and body lengths of 2.8 to 3.1 mm.
The external gills are moderately long, slender, and filamentous; the
yolk sac is still moderately large. The body and anterior part of the
caudal musculature are dark brown; posteriorly the caudal musculature is
pale brown. The caudal fins are creamy tan. The oral discs are large and
ovoid. The growth of the tadpole is summarized in Table 10.

A typical tadpole in stage 30 of development (KU 87652 from 11 km. N
Vista Hermosa, Oaxaca, México) can be described as follows:

Body length 9.5 mm.; tail length 15.5 mm.; total length 25.0 mm.; body
slightly wider than deep; snout rounded laterally, broadly ovoid
dorsally; eyes widely separated, directed dorsolaterally; nostril about
midway between eye and tip of snout; mouth anteroventral; spiracle
sinistral, slightly posterior to midpoint of body and slightly below
midline; anal tube dextral; caudal musculature slender, barely curved
upward distally; dorsal fin not extending onto body, depth of dorsal fin
slightly more than that of ventral fin on mid-length of tail; dorsal
part of body dark brown; ventral surfaces transparent, lacking pigment;
posterior edge of body pale cream-color; caudal musculature creamy white
with interconnected brown spots; caudal fins transparent with small
brown blotches on dorsal fin and posterior half of ventral fin; iris
coppery bronze in life (Fig. 12). Mouth small, median part of upper lip
bare; rest of mouth bordered by single row of bluntly rounded papillae;
lateral fold present; tooth rows 2/3; all tooth-rows approximately equal
in length; second upper row broadly interrupted medially; other rows
complete; upper beak moderately deep, forming broad arch with slender
lateral processes; lower beak slender, broadly V-shaped; both beaks
finely serrate (Fig. 15C).

All tadpoles having fully developed mouth parts have 2/3 tooth rows.
Little variation is noticeable in coloration. In many specimens the
posterior edge of the body is dark brown instead of pale cream-color.
Mottling is rather dense on the caudal fins in all specimens; in some
individuals pigment is concentrated along the anterior one-third of the
lateral groove. In life the body is dark brown with greenish gold flecks
ventrally; the caudal musculature is gray.

In each of two recently metamorphosed young the snout-vent length is
14.0 mm. Coloration of young in life (KU 87653 from 11 km. N Vista
Hermosa, Oaxaca, México): "Dorsum pale tan with dark brown markings.
Thighs orange-yellow; labial stripe white; iris bronze" (Duellman, field
notes, July 10, 1964.)

_Remarks._--Smith (1953:150) named _cyanosticta_ as a subspecies of
_Hyla phaeota_. The differences in cranial characters and certain
external characters between _phaeota_ and _cyanosticta_ indicate that
they are distinct species. Furthermore, a gap of about 350 kilometers
exists between the known geographic ranges of the two kinds.

_Distribution._--_Smilisca cyanosticta_ inhabits wet forests on the
Atlantic slope of southern México and northern Central America from
northern Oaxaca and southern Veracruz through northern Chiapas in México
and into El Petén and northern Alta Verapaz in Guatemala (Fig. 2).
Apparently the range is discontinuous, for in southern México the
species is found in cloud forest at elevations of 830 to 900 meters on
the northern slopes of the Sierra de Juárez. In the Sierra de Los
Tuxtlas in southern Veracruz the species is found in wet forest at
elevations of 300 to 1200 meters, but is absent in the intervening
lowlands characterized by drier forest. In the west forests of northern
Alta Verapaz and El Petén, Guatemala, the species is found at low
elevations.

_Specimens examined._--78, as follows: MEXICO: =Chiapas=: Monte Libano,
MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH
41588. =Oaxaca=: 11 km. N Vista Hermosa, KU 84918-20 (skeletons),
87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH
57199-201; 8 km. S Yetla, KU 87213, UMMZ 124838 (8). =Veracruz=: Coyame,
UMMZ 111459-60; between Coyame and Tebanco, UMMZ 121198; Dos Amates,
UMMZ 121297; between Laguna de Catemaco and Volcán San Martín, UMMZ
121200; Volcán San Martín, UIMNH 35403-4, 35408-12, UMMZ 118163; SE
slope Volcán San Martín, UMMZ 121199, 121295 (2), 121296, 121298.

GUATEMALA: =Alta Verapaz=: Chinajá, KU 55935-7, 55938 (skeleton). =El=

=Petén=: 10 km. NNW Chinajá (Alta Verapaz), KU 55934; Piedras Negras,
CNHM 99006-7, 99227, UIMNH 28853, USNM 111139-41, 111143-7; 8 km. S
Piedras Negras, CNHM 99008; Semicoch, USNM 35907.

    [Illustration: FIG. 2. Map showing locality records for _Smilisca
       cyanosticta_ (triangles) and _Smilisca phaeota_ (circles).]


=Smilisca phaeota= (Cope)

  _Hyla phaeota_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 14
     (9):358, 1862 [Holotype.--USNM 4347 from Turbo, Colombia; J. Cassin
     collector]. Boulenger, Catalogue Batrachia Salientia in British
     Museum, p. 402, Feb. 1, 1882. Werner, Sitzungsb. Akad. Wiss.
     München, 27:215, 1897. Günther, Biologia Centrali-Americana:
     Reptilia and Batrachia, p. 269, Sept. 1901. Nieden, Das Tierreich,
     Amphibia, Anura I, p. 261, June 1923. Dunn, Occas. Papers Boston
     Soc. Nat. Hist., 5:413, Oct. 10, 1931. Gaige, Hartweg, and Stuart,
     Occas. Papers Mus. Zool. Univ. Michigan, 357:4, Oct. 26, 1937.
     Cooper, Copeia, 2:122, June 30, 1944. Breder, Bull. Amer. Mus. Nat.
     Hist., 86(6):416, Aug. 26, 1946. Smith and Taylor, Bull. U. S. Natl.
     Mus., 194:88, June 17, 1948; Univ. Kansas Sci. Bull, 33:364, March
     20, 1950. Taylor, Univ. Kansas Sci. Bull., 35(1):837, July 1, 1952.
     Brattstrom and Howell, Herpetologica, 10:117, Aug. 1, 1954. Goin,
     Herpetologica, 14:120, July 23, 1958. Cochran, Bull. U. S. Natl.
     Mus., 220:57, 1961.

  _Hyla labialis_ Peters, Monats. Konigl. Akad. Wissen. Berlin, p.
     463, 1863 [Holotype.--ZMB 4913 from "region of Bogotá," Colombia];
     Monats. Konigl. Akad. Wissen. Berlin, p. 618, Oct. 16, 1873.
     Boulenger, Catalogue Batrachia and Salientia in British Museum, p.
     397, Feb. 1, 1882.

  _Hyla baudini dolomedes_ Barbour, Occas. Papers Mus. Zool. Univ.
     Michigan, 129:11, Jan. 25, 1923 [Holotype.--MCZ 8539 from Río
     Esnápe, Sambú Valley, Darién, Panamá; Barbour and Brooks
     collectors]. Barbour and Loveridge, Bull. Mus. Comp. Zool. Harvard,
     69:278, June, 1929.

  _Hyla phaeota phaeota_, Smith, Herpetologica, 8:152, Jan. 30, 1953.
     Minton and Smith, Herpetologica, 16:103, June 17, 1960.

  _Smilisca phaeota_, Starrett, Copeia, 4:303, Dec. 30, 1960.

_Diagnosis._--Size large ([M] 65 mm., [F] 78 mm.); skull as long as
wide, lacking frontoparietal fontanelle; large supraorbital flanges
having straight edges and extending posterolaterally; large squamosal
not in contact with maxillary; tarsal fold moderately wide, full length
of tarsus; inner metatarsal tubercle moderately large, low, flat,
elliptical; hind limbs relatively long, tibia averaging more than 53 per
cent of snout-vent length; labial stripe silvery white; lips lacking
vertical bars; loreal region pale green; dark brown or black tympanic
mark dispersing into brown venated pattern on flanks; posterior surfaces
of thighs pale brown, with or without darker flecks or small
cream-colored flecks. (Foregoing combination of characters
distinguishing _S. phaeota_ from any other species in genus.)


  Table 2.--Geographic Variation in Size and Proportions in Males of
     Smilisca phaeota. (Means in Parentheses Below Observed Ranges;
     Data Based on Ten Specimens From Each Locality.)

  ==================================================================
                               |Snout-vent |Head width/|Interorbital
  Locality                     |length     |snout-vent |distance/
                               |           |length     |head width
  -----------------------------+-----------+-----------+------------
  Bonanza, Nicaragua           | 40.8-47.7 | 34.1-38.0 | 31.0-36.1
                               |  (43.7)   |  (36.3)   |  (35.4)
  -----------------------------+-----------+-----------+------------
  Heredia Prov., Costa Rica    | 46.3-59.0 | 32.5-36.0 | 30.5-39.6
                               |  (51.7)   |  (35.0)   |  (34.7)
  -----------------------------+-----------+-----------+------------
  Puntarenas Prov., Costa Rica | 53.6-64.9 | 32.6-36.1 | 31.0-38.0
                               |  (61.4)   |  (34.5)   |  (34.4)
  -----------------------------+-----------+-----------+------------
  Canal Zone, Panamá           | 52.4-65.5 | 33.5-37.6 | 31.3-37.2
                               |  (56.5)   |  (35.6)   |  (34.7)
  -----------------------------+-----------+-----------+------------
  Río Quesada, Colombia        | 52.6-61.0 | 33.1-37.1 | 30.1-33.9
                               |  (56.0)   |  (35.0)   |  (32.1)
  ------------------------------------------------------------------


_Description and Variation._--For the purposes of analyzing geographic
variation in size and proportions, measurements were taken on ten adult
males from each of five samples throughout the range of the species.
Aside from the data summarized in Table 2, the average ratio of tibia
length to snout-vent length is noticeably less in Colombian specimens
(53.4 per cent, as compared with 54.8 to 57.8 per cent in the other
samples) and the ratio of head length to snout-vent length is
noticeably less in Costa Rican specimens (33.5 per cent as compared with
34.9 to 35.1 per cent in the other samples). Also, specimens from
Heredia Province, Costa Rica, have a relatively smaller tympanum (62.7
to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared
with means of 74.0 to 77.9 per cent in the other samples).

Two populations are distinctive as regards the size of adult males.
Specimens from the northern Caribbean lowlands of Nicaragua (Bonanza,
the northernmost locality for the species) are remarkably small. Males
having snout-vent lengths of between 40 and 43 mm. were breeding; the
largest male found had a snout-vent length of 47.7 mm. The other extreme
in size is attained in specimens from the Pacific lowlands of eastern
Costa Rica and western Panamá, where most breeding males have snout-vent
lengths of more than 55 mm.; the largest male had a snout-vent length of
64.9 mm.

The rather striking differences in size among certain samples and the
minor differences in proportions among other samples show no geographic
trends. Instead, the variations apparently are random among the samples.
The data presented here possibly are the results of inadequate sampling,
but more likely reflect actual differences in the populations.

The dorsal ground color of _Smilisca phaeota_ is pale green to tan; the
venter is creamy white. The dorsum is variously marked with dark
olive-green or dark brown spots or blotches (Pl. 6C). A dark
interorbital bar is usually present. Usually a large dark dorsal mark
extends from the occiput to the sacral region, but in many individuals
this blotch is replaced by two or three dark marks. The dorsal markings
are irregular in shape and do not tend to form transverse bands or
longitudinal bars. The hind limbs are marked by dark transverse bands,
usually four or five on the thigh, five or six on the shank, and four on
the tarsus. Two or three narrow bands are usually present on the
proximal part of the fourth toe. The webbing on the feet is brown. The
loreal region is pale green, bordered above by a narrow dark brown
canthal stripe extending from the nostril to the orbit. The upper lip is
silvery white. A broad dark brown or black mark extends posteriorly from
the orbit, encompassing the tympanum, to a point above the insertion of
the forelimb. The flanks are pale green or pale tan and marked with a
fine dark brown or black venation. The anterior surfaces of the thighs
usually are pale brown or grayish tan, sometimes having small,
indistinct darker flecks. The posterior surfaces of the thighs are
similarly colored, but in most specimens small but distinct dark flecks
are present; in some specimens small cream-colored spots are also
present on the posterior surfaces of the thighs. A distinct, narrow
creamy white anal stripe usually is present. A distinct white stripe is
present on the outer edge of the tarsus and fifth toe; on the tarsus the
white stripe is bordered below by dark brown. A white stripe also is
present on the outer edge of the forearm and fourth finger. In breeding
males the throat is dark gray.

Little geographic variation in color or pattern is evident. Few, if any,
specimens from the Pacific lowlands of South America are green in life.
(We have seen no living individuals from South America.) Some living
individuals from Costa Rica and all those seen alive from Nicaragua have
a tint of pale blue on the flanks. In some specimens the dorsal pattern
is so faint as to be barely discernible, whereas in most specimens the
pattern is bold.

The coloration in the living frogs is highly variable due to extreme
metachrosis. Individuals of this species are capable of changing the
dorsal coloration from green to brown in a short period of time. Both
green and brown individuals have been found active at night. Usually
those individuals found hiding by day are brown. One individual from
Finca La Sumbadora, Panamá (now KU 91914), was kept alive in the
laboratory for nearly one month. This individual usually was pale green
with tan dorsal markings at night and tan with pale green markings by
day. On occasion the pale green dorsal markings were boldly outlined by
bright dark green.

In living individuals from throughout the range of the species the iris
is a bronze color, darkest medially with fine black reticulations.

_Natural History._--_Smilisca phaeota_ inhabits humid lowland tropical
forest and seldom ascends the foothills to more than 1,000 meters. The
rather equable climatic conditions, especially more or less evenly
distributed rainfall throughout the year, permit this frog to be active
most of the year. Dunn (1931:413) reported males calling on Barro
Colorado Island, Panamá, in February and in July, and Breder (1946:416)
noted calling individuals in the Chucanaque drainage of Darién, Panamá
in January, March, July, August and October and in Costa Rica in April
through August inclusively. Calling males were found at Bonanza,
Nicaragua in March and in July.

At all times of year the usual daytime retreats for these frogs are near
water; the frogs have been found in elephant ear plants (_Xanthosoma_)
and in bromeliads; occasional individuals have been found sitting on
shaded branches of bushes and trees. None has been observed on the
ground or beneath ground-cover by day.

The length of the breeding season cannot be determined definitely. The
earliest date on which eggs have been found is May 23; Gaige, Hartweg,
and Stuart (1937:5) reported a gravid female taken at El Recreo,
Nicaragua, in September, and we have a gravid female taken at Almirante,
Panamá, in March.

Males usually call from secluded spots at the edge of water. All calling
males that we observed were on the ground within a few centimeters of
the water. The males usually are hidden beneath an overhanging leaf or
some other cover; they definitely do not sit in the open like _Smilisca
baudini_. Most calling males and clasping pairs have been found at the
edges of small pools or shallow ditches, although occasional individuals
are found at the edges of large ponds or streams.

The breeding call consists of one or two moderately short, low-pitched
notes (duration 0.33 to 0.42 seconds), repeated at intervals of about 20
seconds to several minutes. Each note is a low, vibrant "wauk," having
100 to 130 pulses per second and a dominant frequency of 330 to 420
cycles per second (Pl. 10C).

The eggs are deposited in loose clumps amidst vegetation in the water.
Hatchling tadpoles have total lengths of 8.7 to 10.6 mm., and body
lengths of 4.1 to 4.5 mm. The external gills are long and filamentous,
and the yolk sac is large. The head and caudal musculature are dark
brownish black, and the caudal fins are gray. The oral discs are large
and roughly circular. The growth and development of the tadpoles are
summarized in table 11 and figure 16.

A typical tadpole in stage 30 of development (KU 68482 from the Río
Chitaría, Cartago Province, Costa Rica) may be described as follows:
body length 9.7 mm.; tail length 14.6 mm.; total length 24.3 mm.; body
as wide as deep; snout rounded dorsally and laterally; eyes widely
separated, directed dorsolaterally; nostril about midway between eye
and tip of snout; mouth anteroventral; spiracle sinistral, about midway
on length of body and slightly below midline; anal tube dextral; caudal
musculature slender, curved upward distally; dorsal fin extending onto
body; depth of dorsal fin slightly less than that of ventral fin at
mid-length of tail; dorsal part of body pale brown; ventral surfaces
transparent with scattered pigment; pale cream-colored, crescent-shaped
mark on posterior edge of body; caudal musculature pale creamy tan with
scattered pale brown spots; caudal fins transparent with scattered small
brown blotches on dorsal and ventral fins; iris pale bronze in life
(Fig. 13); mouth small; median part of upper lip bare; rest of mouth
bordered by one row of pointed papillae; lateral fold present;
tooth-rows 2/3, first upper row longest; second upper row slightly
shorter, broadly interrupted medially; three lower rows complete, equal
in length, slightly shorter than second upper row; upper beak moderately
deep, forming broad arch with slender lateral processes; lower beak
slender, broadly V-shaped; both beaks serrate (Fig. 15E).

In tadpoles having fully developed mouthparts the tooth-row formula of
2/3 is invariable. The pale crescent-shaped mark on the posterior part
of the body curves anterodorsally on the dorsal surface of the body.
These marks in dorsal view give the appearance of a pair of short, pale
stripes on the posterior part of the body. Most specimens from Costa
Rica have the pale coloration like that described above, but some
individuals (notable KU 87683 from Guápiles, Costa Rica, KU 87707 from
Finca Tepeyac, Nicaragua, and KU 87708 from Bonanza, Nicaragua) have
much more pigment. In these specimens the same color pattern obtains as
in the pallid individuals, but the pigmentation is dense. This is
especially noticeable on the tail.

Recently metamorphosed young have snout-vent lengths of 12.7 to 16.7 mm.
(average, 14.3 mm. in eleven specimens). Coloration of young in life (KU
68484 from Río Chitaría, Cartago Province, Costa Rica): "Dorsum pale
tan; side of head and flanks darker brown, separated from tan dorsum by
an indistinct cream stripe. Limbs pale yellow; thighs flecked with
brown; shank and tarsus yellowish tan with indistinct brown bars. Soles
of feet brown. Belly white; throat dusty cream flecked with silvery
white. Upper lip silvery white. Iris bright gold with black flecks.
Heels, tarsal and anal stripes white" (Duellman, field notes, May 23,
1961).

_Remarks._--Peters (1863:463) named _Hyla labialis_ from the "region of
Bogotá, Colombia", but in 1873 regarded his new species as identical
with _Hyla phaeota_ Cope, 1862, from Turbo, Colombia. The name _Hyla
labialis_ has been used for frogs from the northern Andes in Colombia
(see Dunn, 1944:72, and Stebbins and Hendrickson, 1959:522, for
discussion of nomenclature). Rivero (1961:131) used the name _Hyla
vilsoniana_ Cope, 1899, for the frogs from the northern Andes previously
referred to _Hyla labialis_. A review of the nomenclature and taxonomy
of these frogs, which superficially resemble _Smilisca_ but are
unrelated, is beyond the scope of the present study.

_Hyla baudini dolomedes_ Barbour, 1923, is based on a small _Smilisca
phaeota_ (MCZ 8539) having a snout-vent length of 45.5 mm. Dunn
(1931a:413) placed _dolomedes_ in the synonymy of _Smilisca phaeota_. We
have examined the holotype of _dolomedes_ and agree with Dunn's
assignment.

Smith (1953:150) described _Hyla phaeota cyanosticta_ from Guatemala.
Our studies on the external morphology, coloration, and especially the
cranial osteology provide evidence that _cyanosticta_ is a species
distinct from _phaeota_.

_Distribution._--_Smilisca phaeota_ inhabits humid tropical forests from
northeastern Nicaragua southward on the Caribbean lowlands to elevations
of about 1000 meters and on the Pacific lowlands of Costa Rica,
exclusive of the arid regions of Guanacaste, throughout the lowlands of
Panamá, exclusive of the savannas of the Pacific lowland and the Azuero
Peninsula, and southward on the Pacific slopes of South America through
Colombia to west-central Ecuador; also the valleys of the Río Cauca and
Río Magdalena in Colombia (Fig. 2).

_Specimens examined._--528, as follows: NICARAGUA: =Matagalpa=: Finca
Tepeyac, 10 km. N, 9 km. E Matagalpa, KU 85439, 87707 (tadpoles);
Matagalpa, MCZ 3546-7, UMMZ 92367; 19 km. N Matagalpa, UMMZ 116495-6.
Zelaya: Bonanza, KU 84854-62, 84950-2 (skeletons), 85440-50, 87708-9
(tadpoles); Cukra, AMNH 80618; Río Mico, 16 km. E Recreo, UMMZ 79711
(6), 79712 (4); junction Río Mico and Río Siguia, UMMZ 79713 (10); Río
Siguia, 11 km. NW Rama, UMMZ 79714 (14), 79715 (11), 79716 (21), 79717,
79718 (3).

COSTA RICA: =Alajuela=: Cinchona, KU 32255, 64286-8; 5 km. S Ciudad
Quesada, USC 8077; Laguna Monte Alegre, KU 64289-90; Las Playuelas, 11
km. S Los Chiles, USC 7216; San Carlos, USNM 29961.

=Cartago=: Moravia de Turrialba, KU 32212-47, 37133-5, 41093 (skeleton),
64280-1, USC 7243 (3); Peralta, KU 32271-2; Río Chitaría, 3 km. NNE
Pavones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young);
Río Reventazón, MCZ 29196-203, UMMZ 117677 (9); Turrialba, KU 25720-2,
32209-11, 32266-8, 32273-4, 37136-67, 41090-2 (skeletons), 64270-2, MCZ
29221, 29222 (tadpoles), 29269-70, USNM 29934.

=Guanacaste=: Tilarán, KU 36805-7; 8 km. NE Tilarán, KU 36803-4.

=Heredia=: Barranca del Río Sarapiquí below Isla Bonita, KU 64282-3;
Cariblanco, KU 32256-60, 41094 (skeleton), 64284, MCZ 7967; Isla Bonita,
KU 32250-4; 4.2 km. W Puerto Viejo, KU 64285, 68485; 7.5 km. W Puerto
Viejo, KU 68486; 1 km. S Puerto Viejo, KU 86518.

=Limón=: Bambú, USC 7182 (4); Batán, UMMZ 118582; Coén, MCZ 9825; La
Lola, KU 32262-4, UF 4029, UMMZ 117678 (3); Los Diamantes, CNHM
101295-8, KU 25723-4, 32265, 64267-9; Pandora, UMMZ 122650 (2), USC 7188
(3), 7190; Puerto Limón, KU 32261; Río Larí at Río Dipari, 21 km. SW
Amubre, USC 7177; Río Toro Amarillo, 7 km. W Guápiles, KU 86519, 87683
(tadpoles); Suretka, KU 36808-10, 37168.

=Puntarenas=: Agua Buena, KU 36790; 1.6 km. E Buenos Aires, UMMZ 117578;
3 km. NW Buenos Aires, KU 64304; 4 km. N, 15 km. W Dominical, KU 68491-2
(tadpoles); Esparta, MCZ 8029-30, 8032; Golfito, KU 32270; 6 km. E
Golfito, KU 84999-500 (skeletons); Gromaco, UMMZ 123677 (4); Palmar, KU
32269; 4 km. ESE Palmar Sur, KU 64305-6; 5.6 km. SE Palmar Sur, KU 68489
(tadpoles); 7.0 km. SE Palmar Sur, KU 68490 (young); 8.5 km. SE Piedras
Blancas, KU 64292-303; Quebrada Boruca, 22 km. E Palmar Norte, KU 64291;
Rincón, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), USC 7254; Río
Ferruviosa, 7 km. S Rincón, USC 7235; 1.6 km. WNW Villa Neily, KU 68493
(young), 68494 (tadpoles).

=San José=: San Isidro el General, KU 32249, UMMZ 75025; 10 km. N San
Isidro el General, MCZ 29099-103; 13 km. WSW San Isidro el General, KU
86517; 15 km. WSW San Isidro el General, KU 68487 (tadpoles), 68488
(young), 68495 (young); 20 km. WSW San Isidro el General, KU 32248.

PANAMA: No province: Cano Saddle, USNM 69588; Punta de Pena, USNM 38733;
Quipo, AMNH 18925-6. =Bocas del Toro=: Almirante, KU 80080, 91835-6; 1.6
km. W Almirante, KU 91837; 3 km. W Almirante, KU 91824 (skeleton),
91838-43, 91906-7; 11 km. NW Almirante, CNHM 67853-61; 13 km. W
Almirante, KU 91825-7 (skeletons), 91844-9; Fish Creek, KU 92329; Isla
Popa, KU 91850-1. =Canal Zone=: Barro Colorado Island, CNHM 6007, 13316,
13325, 13331, 13360-2, 13377-8, MCZ 24191-5, UF 7523, UMMZ 63547-60,
64457, 69497 (3); 3.7 km. W Cocoli, KU 67916; Fort Sherman, MCZ 10139;
Gatun, MCZ 35644; Junction roads C25B and C16, TNHC 23839; Madden
Forest Preserve, TNHC 23837-8. =Coclé=: El Valle, KU 77521-4, 77649
(tadpole), TNHC 23369. =Comarca del Barú=: Progreso, UMMZ 61085-9.
Colón: Achiote, KU 77516-20, 77648 (young); Río Candelaria, CNHM
67851-2. =Darién=: Río Esnápe, Sambú Valley, MCZ 8539; Río Sucubti,
Chalichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek,
Camp Townsend, AMNH 40988. =Panamá=: NW slope Cerro Prominente, KU
80459; Finca La Sumbadora, KU 91914 (skeleton). =Chiriquí=: 2 km. W
Concepción, AMNH 68910.

COLUMBIA: =Antioquia=: Puerto Berrio, CNHM 30805 (Goin); Turbo, USNM
39899. =Caldas=: Pueblorrica, Santa Cecilia, CNHM 54768-71 (Goin).
=Choco=: No specific locality, AMNH 3984-6; Andagoya, BMNH 1915.10.21.
69-70, CNHM 81857 (Goin); Golfo de Urabá, CNHM 63881 (Goin); Peña Lisa,
Condoto, BMNH 1913.11.12. 118-125, 1913.11.12. 137-146 (Goin); Pizarro,
CNHM 4451-3, 4455-61 (Goin); Río San Juan, Playa del Oro, CNHM 54772
(Goin); Río Quesada, AMNH 13615-77; 37 km. up Río Puné, AMNH 13688; 48
km. up Río Puné, AMNH 13689. =Narino=: Tumaco, Río Rosario, CJG 2310-13
(Goin). =Valle=: Buenaventura, BMNH 1895.11.16.82 (Goin); Raposa, WAT
166, 346-47, 388 (Goin); Río Calima above Córdoba, CJG 2249-57 (Goin).

ECUADOR: No province: Bulun, AMNH 10620. =Esmeraldas=: Cachabé, AMNH
10625-8; Río Capayas, CNHM 35712; Río Sapaya, UMMZ 58910 (5); Salidero,
AMNH 10623-4; San Javier, AMNH 10618. =Guayas=: Hacienda Balao Chico,
UMMZ 123904. =Imbabura=: Pambelar, AMNH 10629, 10631. =Pichincha=:
Hacienda Espinosa, 9 km. W Santo Domingo de los Colorados, KU 40220.


=Smilisca puma= (Cope), new combination

  _Hyla puma_ Cope, Proc. Amer. Philos. Soc., 22:183, 1885
     [Holotype.--USNM 13735 from Nicaragua; J. F. Moser collector].
     Günther, Biologia Centrali-Americana: Reptilia and Batrachia, p.
     270, Sept., 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 251,
     June, 1923. Cochran, Bull. U. S. Natl. Mus., 220:58, 1961.

  _Hyla wellmanorum_ Taylor, Univ. Kansas Sci. Bull. 25(1):843, July
     1, 1952 [Holotype.--KU 30302 from Batán, Limón, Costa Rica, Edward
     H. Taylor collector]; Univ. Kansas Sci. Bull., 36(1):626, June 1,
     1954. Duellman and Berg, Univ. Kansas Publ. Mus. Nat. Hist.,
     15:194, Oct. 26, 1962.

  _Smilisca wellmanorum_, Starrett, Copeia, 4:303, Dec. 30, 1960.

_Diagnosis._--Size small ([M] 38.0 mm., [F] 46.0 mm.), differing from
other species in the genus by the following combination of characters:
skull about as long as broad; frontoparietal fontanelle keyhole-shaped;
supraorbital flanges absent; squamosal small, not in contact with
maxillary; bony portion of ethmoid terminating at anterior edge of
orbit; tarsal fold weak, two-thirds length of tarsus; inner metatarsal
tubercle small, low, flat, elliptical; snout rounded in dorsal profile;
lips thin and flaring; fingers having only vestige of web; toes one-half
webbed; diameter of tympanum about two-thirds that of eye; narrow labial
stripe white; pair of dark brown (sometimes interconnected) stripes on
tan dorsum; no blue spots on flanks or thighs; vocal sac in breeding
males pale brown. (Foregoing combination of characters distinguishing
_S. puma_ from other species in genus.)

_Description and variation._--Ten breeding males from the vicinity of
Puerto Viejo, Heredia Province, Costa Rica, have snout-vent lengths of
32.5 to 37.9 mm. (34.8 mm.). In these specimens, the length of the tibia
to the snout-vent length is 0.48 to 0.53 (0.51), and the tympanum/eye
ratio is 0.52 to 0.72 (0.65). Seven females have snout-vent lengths of
40.8 to 45.8 mm. (43.9 mm.). No individual has more than a vestige of a
web between the second and third and fourth fingers. None has a web
between the first and second fingers. Breeding males lack nuptial
excrescences on the thumbs. The vocal sac is moderately large and
bilobed.

In preserved specimens the dorsal ground color varies from yellowish tan
to grayish brown. All specimens have dark brown dorsal markings in the
form of a pair of dorsal stripes, variously modified (Pl. 7A). In some
specimens, such as KU 91716, the stripes are discrete and extend from
the postorbital region nearly to the vent. In most specimens the stripes
are connected by a transverse mark in the scapular region and in many
others also by a transverse mark in the sacral region. In some specimens
the stripes are fragmented posteriorly; fragmentation is extreme in KU
30300, in which the dorsal pattern consists of two series of dark
longitudinal dashes. The other extreme is a nearly complete fusion of
the stripes, as in KU 91714. A dark brown interorbital bar usually
extends onto the eyelids, but in some specimens this is reduced to a
short V-shaped mark or small spot between the eyes. There is no dark
post-tympanic mark, but dark brown pigment forms a venated pattern from
the axilla to the mid-flank; the inguinal region is white, finely
mottled with dark brown. The dorsal surfaces of the hind limbs are
colored like the body and have two or three dark brown transverse marks
on the thighs, three to five marks on the shanks, and one or two marks
or irregularly arranged dark flecks on the tarsi. The anterior and
posterior surfaces of the thighs are pale tan to brown. The webbing of
the feet is tan to grayish brown. A narrow white labial stripe, white
anal stripe, and narrow white stripes on the tarsi and outer edges of
the forelimbs are invariably present. The ventral surfaces are creamy
white.

In life the dorsum is tan or pale brown with dark brown markings. Some
individuals have scattered metallic green flecks on the dorsum. The
flanks are mottled dark brown and creamy white. The posterior surfaces
of the thighs are dark brown. The vocal sacs are grayish brown, and the
iris is a deep bronze color.

_Natural History._--_Smilisca puma_ inhabits humid lowland tropical
forests having more or less evenly distributed rainfall throughout the
year. The equable climatic conditions seemingly permit these frogs to be
active throughout most of the year. Taylor (1952:846) found calling
males at Batán, Costa Rica, on July 20, 1951. We found the species
breeding near Puerto Viejo, Costa Rica, on February 19, June 18, July
13, and July 31. Specimens of calling males from Costa Rica in the
collection at the University of Southern California were obtained in
February at La Fortuna, on August 22 at Los Diamantes, on August 30 at
Jabillos, and on September 5 at La Lola. Gravid females were collected
in June, July and August.

Males call from shallow water. All breeding congregations of this
species that we have found were in a grassy marsh, 7.5 kilometers west
of Puerto Viejo, Costa Rica. Tadpoles were found in water-filled
depressions in the marsh at night. When first observed, tadpoles were
near the surface of the water; they responded to light by quickly taking
refuge in the dense grass. No tadpoles were observed by day.

The breeding call consists of a low squawk, usually followed by a series
of one or more rattling secondary notes (duration of primary notes,
0.06-0.35 seconds; of secondary notes, 0.10 to 0.47 seconds), repeated
at intervals of 5 to 55 seconds. The primary notes have 187 to 240
pulses per second and major frequencies of about 740 to 1870 cycles per
second (Pl. 11A).

Only six tadpoles are available for study. Four of them in stage 34 of
development have body lengths of 9.0 to 9.5 mm., tail lengths of 14.0 to
15.0 mm., and total lengths of 23.0 to 24.5 mm. One tadpole in stage 38
and one in stage 40 have total lengths of 31.0 mm. A typical tadpole in
stage 34 of development (KU 91807 from 7.5 km. W Puerto Viejo, Heredia
Province, Costa Rica) has a body length of 9.5 mm., tail length of 15.0
mm., and total length of 24.5 mm.; body about three-fourths as deep as
wide; snout rounded dorsally and laterally; eyes widely separated,
directed dorsolaterally; nostril about midway between eye and tip of
snout; mouth anteroventral; spiracle sinistral, about two-thirds
distance from snout to posterior end of body and slightly below midline;
anal tube dextral; caudal musculature slender, barely curved upward
distally; dorsal fin extending onto body; at mid-length of tail, depth
of caudal musculature equal to that of dorsal fin and ventral fin; body
grayish brown, palest ventrally; caudal musculature pale creamy yellow
with bold gray reticulations; caudal fins transparent with gray
reticulations anteriorly and black flecks posteriorly on both fins (Fig.
14A). Median part of upper lip bare; rest of mouth bordered by two rows
of short blunt papillae; lateral fold present; tooth-rows 2/3; upper
rows equal in length; second upper row broadly interrupted medially;
three lower rows complete, first and second rows equal in length,
slightly shorter than upper rows; third lower row noticeably shorter;
upper beak shallow, forming broad, continuous arch with slender lateral
processes; lower beak slender, broadly V-shaped, both beaks finely
serrate (Fig. 15B).

All six tadpoles are colored alike, except that in the larger specimens
scattered white flecks are present on the ventral surface of the body,
and the dark reticulations continue farther posteriorly on the caudal
fins than in the smaller tadpoles. In two specimens the third lower
tooth-row is only about one-half the length of the other lower rows, and
in one specimen the second lower tooth-row is shorter than the first.
Coloration of tadpoles in life: "Body olive-brown with silvery green
flecks laterally. Caudal musculature olive-brown with greenish tan
flecks. Fins brown with greenish gold flecks. Iris deep bronze."
(Duellman, field notes, February 19, 1965).

One recently metamorphosed young (KU 91808) has a snout-vent length of
12.4 mm. In life this frog had a pale tan dorsum with dark brown
markings, yellowish tan posterior surfaces of thighs, grayish brown
throat, and bronze iris.

_Remarks._--The identity of Cope's _Hyla puma_ has not been known. The
name has appeared in various compilations, but no workers have referred
any of their specimens to that species. Examination of the holotype
(USNM 13735), an adult female, revealed the presence of the following
combination of characters: snout-vent length 45.8 mm., snout blunt above
and rounded laterally, nostrils close to tip of snout, lips thin and
flaring, a vestige of a web on the hands, feet about one-half webbed,
tarsal fold weak and extending about two-thirds length of tarsus, dorsal
markings consisting of a faded dark interorbital bar and a pair of faded
longitudinal brown marks connected by a transverse band in the scapular
region. The type agrees well with specimens of _Smilisca wellmanorum_
(Taylor, 1952); the vestigial webbing on the hands and the dorsal
coloration are especially significant. Consequently, we consider _Hyla
wellmanorum_ Taylor, 1952, to be a synonym of _Hyla puma_ Cope, 1885.
Cope gave only "Nicaragua" as the locality for _Hyla puma_. The specimen
was part of a collection received at the United States National Museum
from Lt. J. F. Moser. Among the species in the collection are
_Dentrobates pumilio_, _Phyllomedusa helenae_, _Corythophanes
cristatus_, _Pliocercus dimidatus_, _Tretanorhinus nigroluteus_, and
others characteristically found on the Caribbean lowlands of Central
America. Thus, it seems reasonable to assume that the type specimen of
_Hyla puma_ came from the Caribbean lowlands. Though no other Nicaraguan
specimens have been found by us, numerous specimens are known from the
Caribbean lowlands of Costa Rica.

Cochran (1961:58), in her catalogue of type specimens in the United
States National Museum, listed _Hyla puma_ Cope, 1885, as a synonym of
_Hyla molitor_ Schmidt, 1857. She made no qualifying statements. Schmidt
(1858:246), in his descriptions of the species in the year following his
publication of the names and Latin diagnoses, stated: "Dorsum uniformly
gray, more intensive on back, fading away laterally and on extremities;
in every-day-life this blue would be called _Mueller's Blau_. A
delicately dotted black line runs on the canthus rostralis from the
opening of the nose to the corner of the eye. In the armpits, on the
flanks and the thighs two of our three specimens have black marblings."
[Free translation] Certainly on the basis of coloration _Hyla puma_ is
distinctly different from _Hyla molitor_.

_Distribution._--This species lives in the wet, forested regions of the
Caribbean lowlands of Costa Rica and presumably southern Nicaragua (Fig.
3). All specimens are from low elevations; the highest known elevation
for the occurrence of this frog is 285 meters at Laguna Bonilla.

    [Illustration: FIG. 3. Map showing locality records for _Smilisca
       puma_ (triangles) and _Smilisca sila_ (circles).]

_Specimens examined._--62, as follows: NICARAGUA: No specific locality,
USNM 13735.

COSTA RICA: =Alajuela=: Jabillos, 5 km. N Santa Clara, USC 8058 (6); 5
km. W La Fortuna, USC 8078 (2); Río La Fortuna at La Fortuna, USC 7151
(3). =Cartago=: Laguna Bonilla, tunnel camp near Peralta, KU 32171.
=Heredia=: Puerto Viejo, KU 86521; 5.9 km. W Puerto Viejo, KU 64307; 7.5
km. W Puerto Viejo, KU 64308-10, 64311 (skeleton), 64312-15, 68635-6
(skeletons), 85001-2 (skeletons), 86520, 87770-1 (skeletons), 91709-16,
91791-2, 91807 (tadpoles), 91808 (young). =Limon=: Batán, KU 30300-2; La
Lola, KU 32169, USC 141, 201, 8067; Los Diamantes, KU 32170, UMMZ 118470
(6), USC 212; 2.4 km. E Los Diamantes, USC 8049 (5).


=Smilisca sila= new species

  _Hyla gabbi_, Noble, Proc. Biol. Soc. Washington, 37:66, Feb. 21,
     1924. Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, Oct. 10,
     1931. Schmidt, Smithsonian Misc. Coll., 89(1):6, March 16, 1933.

   _Hyla sordida_, Dunn, Copeia, 3:166, Nov. 19, 1937. Cooper, Copeia,
     2:121, June 30, 1944. Breder, Bull. Amer. Mus. Nat. Hist.,
     86(8):417, Aug. 26, 1946.

  _Hyla phaeota_, Breder, Bull. Amer. Mus. Nat. Hist., 86(8): pl. 55,
     Aug. 26, 1946.

_Holotype._--Adult male, KU 91852 from a small stream at the north edge
of the village of El Volcán, Chiriquí Province, Panamá, elevation 1280
meters; obtained on Feb. 5, 1965, by William E. Duellman.

_Paratypes._--KU 91853-74, collected with the holotype.

_Diagnosis._--Size moderate ([M] 45.0 mm., [F] 62.2 mm.); skull wider
than long, having large, ovoid frontoparietal fontanelle; supraorbital
flanges absent; squamosal small, not contacting maxillary; bony section
of ethmoid extending anteriorly between nasals; tarsal fold weak, full
length of tarsus; inner metatarsal tubercle low, flat, elliptical; lips
thick, rounded, not flaring; fingers one-third webbed; toes
three-fourths webbed; diameter of tympanum about one-half that of eye;
margin of upper lip faintly marked by interrupted creamy white stripe;
dark spots on dorsum; pale flecks on flanks and posterior surfaces of
thighs; vocal sacs in breeding males dark brown. (Foregoing combination
of characters distinguishing _S. sila_ from any other species in genus.)

_Description of holotype._--Snout-vent length 36.6 mm.; tibia length
19.8 mm., 54.1 per cent of snout-vent length; foot length 15.5 mm., 42.3
per cent of snout-vent length; head length 12.7 mm., 34.7 per cent of
snout-vent length; head width 13.3 mm., 36.8 per cent of snout-vent
length; snout short, in lateral profile truncate, only slightly rounded
above, in dorsal profile rounded; canthus rounded; loreal region
noticeably concave; lips thick, rounded, not flaring; nostrils not
protuberant, directed laterally; internarial distance 3.0 mm.;
internarial area flat; top of head flat; interorbital distance 3.5 mm.,
26.3 per cent of head width; diameter of eye 4.2 mm., thrice distance
(1.4 mm.) from tympanum to eye, and half again distance (2.8 mm.) from
orbit to nostril; pupil horizontally ovoid; width of eyelid 2.8 mm.,
21.1 per cent of head width; dermal fold from posterior corner of orbit
covering upper edge of tympanum to point above insertion of forelimb;
diameter of tympanum 2.3 mm., 54.7 per cent of diameter of eye; no
axillary membrane; arms moderately robust; weak fold on wrist; faintly
scalloped fold along ventrolateral margin of forearm; fingers short,
slender; fingers from shortest to longest, 1-2-4-3; vestige of web
between first and second fingers; others about two-fifths webbed; discs
moderate, diameter of that on third finger about one-third diameter of
eye; triangular outer palmar tubercle; elliptical inner palmar tubercle
on base of pollex; subarticular tubercles large, conical, none bifid;
supernumerary tubercles few, large, conical; brown nuptial excrescence
on prepollex; heels overlap by about one-fifth length of shank when hind
limbs adpressed; tibiotarsal articulation extending to nostril; tarsal
fold weak, extending nearly full length of tarsus; inner metatarsal
tubercle elliptical, flat; outer metatarsal tubercle absent; toes
moderately long; toes from shortest to longest, 1-2-3-5-4, third and
fifth about equal in length; discs about same size as those on fingers;
webbing extending to middle of penultimate phalanx on all toes, except
only to distal end of antepenultimate phalanx of fourth toe;
subarticular tubercles round; supernumerary tubercles large, round,
present only on proximal digits. Anal opening directed posteriorly at
level of upper edge of thighs; no noticeable anal sheath; flat tubercles
ventrolateral to anal opening large; skin of chest, belly, and posterior
surfaces of thighs granular; other surfaces smooth; tongue broadly
cordiform, shallowly notched posteriorly, and barely free behind;
vomerine teeth 4-4, situated on ventral surfaces of separated rounded
prominences between posterior margins of small, ovoid inner nares; vocal
slits long, each situated along inner margin of ramus; color (in
preservative) pinkish tan above with irregular olive-brown markings
forming interconnected spots on back; four bars on dorsal surface of
each thigh; five bars on shank, and three on tarsus; inguinal region
white with black mottling; posterior surfaces of thighs yellowish tan
proximally, dark brown distally; margins of lips grayish white with
brown markings; ventral surfaces of hands and feet grayish brown; belly
and posterior part of throat creamy white; anterior part of throat
brown.

_Description and variation._--Ten breeding males from Finca La
Sumbadora, Panamá, have snout-vent lengths of 40.0 to 44.8 mm. (42.3
mm.). In these specimens the tibia/snout-vent length ratio is 0.50 to
0.57 (0.54), and the tympanum/eye ratio is 0.48 to 0.58 (0.53). There is
a geographic gradient in size; specimens from the western part of the
range (southern Costa Rica) are smaller than those from the eastern part
of the range (eastern Panamá). Five males from the Pacific lowlands of
southern Costa Rica have snout-vent lengths of 31.6 to 38.2 mm. (34.7
mm.); ten males from El Volcán, Chiriquí, Panamá, 32.6 to 37.9 mm. (36.4
mm.), and eight males from Barro Colorado Island, Canal Zone, 38.2 to
42.0 mm. (35.6 mm.). These are smaller than the males from Finca La
Sumbadora, which is east of the Canal Zone. Ten females from El Volcán
have snout-vent lengths of 44.2 to 55.6 mm. (49.2 mm.), as compared 56.1
to 62.2 mm. (58.2 mm.) in three females from Finca La Sumbadora.

Large females have scattered small tubercles on the head and back;
tubercles occur in males from Costa Rica and in some males from western
Panamá. The truncate snout is characteristic of both sexes.

The coloration of _Smilisca sila_ consists of a gray, tan, or pale
reddish brown dorsal ground color and a creamy white venter. The dorsum
is marked by dark brown, olive-brown, or dark reddish brown spots or
blotches (Pl. 7B). Usually the blotches are discrete, but in some
individuals they are interconnected and form an irregular dark mark on
the dorsum. There is no tendency for the blotches to form transverse
bars as in _Smilisca sordida_. In one specimen (KU 80467) the blotches
are fused and form two wide irregular longitudinal stripes, as in
_Smilisca puma_. In some females the dorsal markings are reduced to a
few small spots or are nearly absent (KU 92332), whereas in other
females the dorsal markings are bold. In one female (KU 91894) the
dorsal markings are narrowly bordered by pale blue, and numerous pale
blue flecks are present on the pale brown dorsum. In many individuals of
both sexes small white flecks are present on the dorsal surfaces.

Usually the flanks and posterior surfaces of the thighs have black
mottling enclosing pale blue spots and flecks, respectively. The dorsal
surfaces of the limbs are marked by dark brown transverse bars; usually
three or four bars are present on each forearm, thigh, and shank. The
coloration of the flanks and limbs varies geographically. Specimens from
southern Costa Rica and western Panamá have distinct bars on the limbs;
the posterior surfaces of the thighs have brown reticulations enclosing
small blue flecks in specimens from Costa Rica and bolder, black
reticulations enclosing large pale blue spots in specimens from western
Panamá. In specimens from Costa Rica the flanks are brown with pale blue
flecks, whereas in those from Chiriquí, Panamá, the flanks are pale blue
with dark brown mottling in the inguinal region. Frogs from El Valle and
Cerro la Campana usually have distinct bars on the limbs; the posterior
surfaces of the thighs are colored as in frogs from Chiriquí, and the
inguinal region is pale blue with coarse brown mottling. Specimens from
Barro Colorado Island are marked like those from El Valle and Cerro la
Campana, except that on the posterior surfaces of the thighs fine black
reticulations enclose many dark blue spots. In specimens from Darién and
from Panamá Province east of the Canal Zone (Altos de Pacora, Cerro
Jefe, Finca La Sumbadora, and Río Pacora), the markings on the dorsal
surfaces of the limbs are indistinct or absent in males, but distinct in
some females. Intense brown and black pigment forms fine reticulations
delimiting bold blue spots on the flanks; this coloration extends to the
axilla in many specimens. Fine black reticulations enclose many dark
blue spots on the posterior surfaces of the thighs.

In females, the throat is creamy white; in some specimens scattered
brown flecks are present on the chin and throat. In breeding males the
anterior part of the throat is dark gray or dark brown.

The coloration in life is as variable as it is in preservative. In life
the holotype had a tan dorsum with dark olive-green irregular markings
and small green flecks. The limbs were tan with dark brown transverse
bars. The flanks were grayish tan anteriorly; the inguinal region and
posterior surfaces of thighs were blue with black mottling. The belly
was creamy white, and the throat was brown with creamy yellow flecks.
The iris was a dull bronze color. Among the paratypes, some individuals
had green flecks, others did not. The inguinal region and posterior
surfaces of the thighs were pale blue, pale yellowish green, or grayish
tan with black mottling. The blue was most noticeable in females.

Colors of a male from Finca La Sumbadora, Panamá, were described as
follows: "Dorsum olive-brown; irregular dark brown blotches, pale green
flecks, and raised creamy yellow spots on dorsal surfaces; belly creamy
white; throat grayish brown; undersides of limbs grayish tan; groin,
anterior and posterior surface of thigh, inner surface of shank,
anterior edge of tarsus, and proximal parts of third and fourth toes
pale blue marbled with dark brown and black; webbing brown; iris pale
bronze, finely reticulated with black." (Duellman, field notes, January
28, 1964.)

A female (now KU 91894) from Altos de Pacora, Panamá, was described as
follows: "An irregular dark brown, green-bordered figure on head and
back; dark brown, green-bordered bands on limbs--all on a lighter brown
and heavily green-spotted background. These markings are more vivid at
night than during the day. Lower sides, from midbody onto front of
thighs and rear of thighs onto venter of shanks to heels and thence
dorsally onto basal portions of toes heavily blue spotted on a light
brown (front of thighs and venter of shanks) to blackish brown
background. Venter cream. Iris gray-brown, finely veined with dark
brown." (Charles W. Myers, field notes, December 14, 1964.) Note that in
the earlier discussion of coloration of preserved specimens, the green
spots and borders have changed to pale blue after six months in
alcohol.

In living individuals from Costa Rica and Panamá west of the Canal Zone,
the blue coloration on the flanks and thighs is much less conspicuous
than in specimens from eastern Panamá. The color of the iris is
variable, even in frogs from one locality. The coloration of the iris in
13 living frogs (now KU 92333-45) from Valle Hornito, Chiriquí, Panamá,
was described as follows: "Iris variable--from pale to dark brown; in a
few the iris has a golden cast to the brown; in a few others the lower
half of the iris is pale gray with the upper half being light brown."
(Charles W. Myers, field notes, April 24, 1965).

_Natural history._--_Smilisca sila_ inhabits the Pacific slopes of lower
Central America where a pronounced dry season occurs. We have records of
males calling in December through May and also in August (latter date
from El Volcán, Chiriquí, Panamá). The breeding season seems to be
correlated with the time of the year when the water is clear and at a
low level in the streams where these frogs breed.

Males call from the edges of small, shallow streams, from rocks in the
streams, or less frequently from vegetation overhanging the streams.
Females are most frequently found on the banks of streams, and clasping
pairs usually are in shallow pools in streams. One individual was found
in a bromeliad about three meters above the ground in the daytime.

The breeding call consists of a low squawk, usually followed by a series
of one or more rattling secondary notes (duration of primary notes, 0.06
to 0.28 seconds; of secondary notes, 0.14 to 0.48 seconds), repeated at
intervals of 4 to 20 seconds. The primary notes have 97 to 120 pulses
per second and major frequencies of about 900 to 2220 cycles per second
(Pl. 11B).

Eggs were obtained artificially in the field; the average length of ten
embryos in the neural groove stage is 2.4 mm., and the average diameter
of the outer envelope is 4.9 mm. Hatchlings have large, conical oral
discs, heavy gills, and a large amount of yolk; their average total
length is 6.3 mm.

Tadpoles have been found in pools in clear streams; some tadpoles have
been observed to cling by their mouths to rocks in the stream; others
were found on the bottom where they seek refuge among pebbles or under
rocks and leaves. A complete developmental series of tadpoles is not
available. Eleven tadpoles in stage 25 of development have body lengths
of 8.3 to 10.2 mm. (9.3 mm.), tail lengths of 17.3 to 21.0 mm. (18.8
mm.), and total lengths of 25.9 to 31.0 mm. (28.1 mm.). One tadpole in
stage 41 and one in stage 42 have body lengths of 11.5 and 12.5 mm.,
tail lengths of 27.2 and 29.5 mm., and total lengths of 38.7 and 42.0
mm., respectively. The snout-vent lengths of two specimens in stage 43
and one in stage 45 are 12.7, 13.0, and 13.6 mm., respectively.

A typical tadpole in stage 25 of development (KU 80620 from Finca La
Sumbadora, Panamá) has a body length of 9.5 mm., tail length of 19.0
mm., and a total length of 28.5 mm.; body only slightly wider than deep,
nearly flat dorsally; snout broadly rounded in dorsal view, bluntly
rounded in lateral view; eyes widely separated, directed dorsolaterally;
nostril slightly closer to eye than to tip of snout; mouth ventral;
spiracle sinistral, located about two-thirds distance from snout to
posterior edge of body; anal tube dextral; caudal musculature moderately
heavy, straight; dorsal fin not extending onto body; fins deepest at
about two-fifths length of tail, where depth of caudal musculature about
equal to depth of dorsal and depth of ventral fin; musculature
extending nearly to tip of tail; body dark grayish brown above and pale
grayish tan below with small dark brown spots dorsally and white flecks
laterally; caudal musculature pale tan with dark brown flecks over
entire surface and dark brown streaks on posterior one-half of ventral
fin and on all of dorsal fin (Fig. 14B). Median one-third of upper lip
bare; rest of mouth bordered by a single row of conical papillae;
lateral fold present; tooth rows 2/3; upper rows cone-shaped, about
equal in length, broadly /\-shaped; second upper row narrowly
interrupted medially; lower rows complete, about equal in length, but
slightly shorter than upper rows; upper beak moderately massive, its
inner surface forming a continuous arch with short lateral processes;
lower beak broadly \/-shaped; both beaks finely serrate (Fig. 15D).

Tadpoles from El Volcán, Chiriquí (KU 91833), are more heavily pigmented
than those from Finca La Sombadora; the spots on the tail are larger. In
life these tadpoles had dark brownish black bodies with golden and green
lichenous flecks; the tail was tan with dark brown markings, and the
iris was a grayish bronze color. In life tadpoles from Finca La
Sumbadora were olive-tan above and dark gray with pale bluish gray
irridescent spots ventrally. The caudal musculature was creamy tan with
brown flecks and streaks, and the iris was pale bronze.

Metamorphosing young have been found on vegetation at the edge of
streams and have been raised in the laboratory. Seven recently
metamorphosed young have snout-vent lengths of 13.6 to 15.6 mm. (14.6
mm.). A living juvenile (KU 91913) raised in the laboratory from a
tadpole obtained at Finca La Sumbadora had a brown dorsum with darker
brown markings, a white spot below the eye, and a narrow white labial
stripe. The belly was white; the flanks were brown with white spots, and
the posterior surfaces of the thighs were yellowish tan. The iris was a
golden bronze color with much black reticulation.

_Remarks._--This species has been confused with _Smilisca sordida_; most
authors have referred both species to _Hyla (Smilisca) gabbi_.
Examination of the types of _Hyla sordida_, _gabbi_, _salvini_, and
_nigripes_ revealed that all of the names were referable to a single
species (_S. sordida_), and that the small, blunt-snouted species in
Panamá and southern Costa Rica probably was without a name. Possibly
_Hyla molitor_ Schmidt (1857) is based on the species that we have named
_S. sila_, but several discrepancies in his description, plus the
unknown provenance of the type, have led us to discount the
applicability of that name to the species under consideration.

_Distribution._--_Smilisca sila_ ranges along the Pacific slopes and
lowlands of southern Costa Rica and Panamá at elevations from sea level
to about 1300 meters; in northern South America the species occurs in
the Caribbean lowlands and in the valleys of the northward draining
rivers of Colombia (Fig. 3).

_Specimens examined_, 234, as follows: COSTA RICA: =Puntarenas=: 6 km. E
Golfito, KU 91717; Quebrada Boruca, 22 km. E Palmar Norte, KU 64265-6;
Río Zapote, 7 km. E Palmar Norte, USC 7100 (2). =San José=: San Isidro
el General, KU 28200; 14 km. NW San Isidro el General, USC 7098 (2); 15
km. WSW San Isidro el General, USC 7097.

PANAMA: =Canal Zone=: Barro Colorado Island, AMNH 62320-3, CNHM 13324,
13326-8, 13330, 13338, 13359, 13423-5, KU 80460-6, 80619 (young), 80625
(skeleton), UMMZ 63542-6, USC 7051. =Chiriquí=: Boquete, AMNH 69815,
UMMZ 58441-5; El Volcán, KU 77413, 91828-31 (skeletons), 91852-74, 91832
(eggs), 91833 (tadpoles); 6 km. S El Volcán, CNHM 60442; 16 km. NNW El
Volcán, KU 91879-90; Finca Palosanto, 6 km. WNW El Volcán, KU 77406-12,
77692 (skeleton), 91875-7, 92330-1; Río Colorado, 17 km. NNW El Volcán,
KU 91878, 92332; Valle Hornito, 19 km. NE Gualaca, KU 92333-45. =Coclé=:
El Valle, AMNH 55440-5 (13), 59607-14, CNHM 48140, 60349-2, 60387-92,
60401-4, 60443, 67842-5, KU 91834 (young), 91902-4, TNHC 23751-2, USNM
140653. =Colón=: Río Candelaria, AMNH 53708-15, CNHM 67826-36. =Darién=:
Camp Creek, Camp Townsend, AMNH 40756-7, 40936-9, 40992; Río Chico, AMNH
39784, 40986-7; Río Pita, CNHM 67823-5; Tacarcuna, USNM 141796-802;
Three Falls Creek, AMNH 41684, 51788. =Los Santos=: Cerro Hoya, USNM
148213-4; Lajamina, Río Puria, KU 67915. =Panamá=: Altos de Pacora, KU
91894; Cerro Jefe, KU 91895-6; Cerro La Campana, CNHM 67846, KU
91897-900, USNM 139689; Finca La Sumbadora, KU 80467-81, 80620
(tadpoles), 91910 (eggs), 91911-2 (tadpoles), 91913 (young), 91908-9
(skeletons); Río Calobra, USNM 53722, Río Pacora, 9 km. NNE Pacora, KU
91901. =Veraguas=: Cerro Carbunco, USNM 129066; Cerro Tute, CNHM
67837-41; Isla Cebaco, Río Platanal, KU 91891-3.

COLOMBIA: =Antioquia=: Urabá, Villa Arteaga, CNHM 63893 (Goin).
=Atlantico=: Sabanalarga, Río Causa, AMNH 14506.


=Smilisca sordida= (Peters), new combination

  _Hyla sordida_ Peters, Monatsb. Konigl. Akad. Wissen. Berlin., p.
     460, 1863 [Syntypes.--ZMB 3141 (two specimens) from "Veragua,"
     Panamá; J. von Warszewicz collector]. Brocchi, Mission scientifique
     au Mexique ..., pt. 3, sec. 2, Études sur les batrachiens, p. 42,
     1881. Boulenger, Catalogue Batrachia Salientia in British Museum,
     p. 393, Feb. 1, 1882. Günther, Biologia Centrali-Americana:
     Reptilia and Batrachia, p. 273, Sept. 1901. Nieden, Das Tierreich,
     Amphibia, Anura, I, p. 258, June, 1923.

  _Hyla gabbi_ Cope, Jour. Acad. Nat. Sci. Philadelphia, new ser., 8,
     pt. 2:103, 1876 [Syntypes.--USNM 30658-9 from near Sipurio, Limón,
     Costa Rica; William M. Gabb collector]. Brocchi, Mission
     scientifique au Mexique ..., pt. 3, sec. 2, Études sur les
     batrachiens, p. 37, 1881. Boulenger, Catalogue Batrachia Salientia
     in British Museum, p. 372, Feb. 1, 1882. Cope, Bull. U. S. Natl.
     Mus., 32:32, 1887. Günther, Biologia Centrali-Americana: Reptilia
     and Batrachia, p. 274, Sept. 1901. Werner, Abhand. Konigl. Akad.
     Wissen. München., 22:351, 1903. Nieden, Das Tierreich, Amphibia,
     Anura I, p. 252, June, 1923. Taylor, Univ. Kansas Sci. Bull.,
     35(1):840, July 1, 1952. Cochran, Bull. U. S. Natl. Mus., 220:54,
     1961.

  _Hyla nigripes_ Cope, Jour. Acad. Nat. Sci. Philadelphia, new ser.,
     8, pt. 2:104, 1876 [Syntypes.--USNM 30685-6, from Pico Blanco,
     Costa Rica; William M. Gabb collector]. Brocchi, Mission
     scientifique au Mexique ..., pt. 3, sec. 2, Études sur les
     Batrachiens, p. 38, 1881. Boulenger, Catalogue Batrachia Salientia
     in British Museum, p. 394, Feb. 1, 1882. Cope, Bull. U. S. Natl.
     Mus., 32:32, 1887. Günther, Biologia Centrali-Americana: Reptilia
     and Batrachia, p. 278, Sept., 1901. Nieden, Das Tierreich,
     Amphibia, Anura I, p. 253, June, 1923. James, Copeia, 3:147, Sept.
     30, 1944. Taylor, Univ. Kansas Sci. Bull, 35(1):853, July 1, 1952.
     Cochran, Bull. U. S. Natl. Mus., 220:56, 1961.

  _Hyla salvini_ Boulenger, Catalogue Batrachia Salientia in British
     Museum, p. 372, Feb. 1, 1882 [Syntypes.--BMNH 1947.2.24.13-14 from
     Cartago, Costa Rica; Osbert Salvin collector]. Günther, Biologia
     Centrali-Americana: Reptilia and Batrachia, pl. 71, Fig. B., Sept.,
     1901. Werner, Abhand. Zool.-Bot. Gesell. Wien, 46:8, Sept. 30,
     1896.

  _Smilisca gabbi_, Starrett, Copeia, 4:303, Dec. 30, 1960.

_Diagnosis._--Size moderate ([M] 45 mm., [F] 64 mm.); skull slightly
wider than long, having large and elongate frontoparietal fontanelle;
supraorbital flanges absent; squamosal small, not contacting maxillary;
bony section of ethmoid terminating just anterior to anterior edge of
orbit; tarsal fold weak, full length of tarsus; inner metatarsal
tubercle long, low, flat, elliptical; lips thin and flaring; fingers
one-half webbed; toes four-fifths webbed; diameter of tympanum about
one-half that of eye; no white labial stripe; dorsal dark markings
irregular, sometimes forming broad transverse bars; pale flecks on
flanks and usually on posterior surfaces of thighs; vocal sacs in
breeding males white. (Foregoing combination of characters
distinguishing _S. sordida_ from any other species in genus.)

_Description and variation._--Ten breeding males from 15 to 20
kilometers west-southwest of San Isidro el General, San José, Costa
Rica, have snout-vent lengths of 38.1 to 42.6 mm. (40.5 mm.). In these
specimens, the tibia/snout-vent length ratio is 0.50 to 0.54 (0.52), and
the tympanum/eye ratio is 0.45 to 0.57 (0.49). Specimens from the
Pacific slopes of Costa Rica are larger than those from the Meseta
Central and the Caribbean lowlands. Ten males from 6 kilometers east of
Golfito, Puntarenas, have snout-vent lengths of 38.4 to 44.6 mm. (41.8
mm.), and five males from Rincón, Peninsula de Osa, have snout-vent
lengths of 38.8 to 41.6 mm. (40.3 mm.). Snout-vent lengths of ten males
from La Fortuna, Alajuela, are 31.9 to 36.0 mm. (34.4 mm.), of ten males
from Pandora, Limón, 33.8 to 37.6 mm. (35.9 mm.), and of ten males from
Escazú and Río Jorco on the Meseta Central, 34.3 to 37.6 mm. (36.0 mm.).
Eight females from the Río Jorco on the Meseta Central have snout-vent
lengths of 48.8 to 53.8 mm. (50.4 mm.), and six females from various
localities on the Pacific slopes of Costa Rica have snout-vent lengths
of 56.5 to 64.0 mm. (59.8 mm.). The only noticeable differences in
proportions between males and females is in the tympanum/eye ratio; for
example, this ratio is 0.47 to 0.53 (0.49) and 0.54 to 0.68 (0.61) in
ten males and eight females, respectively, from the Meseta Central.

The shape of the snout and the associated cranial elements of _S.
sordida_ vary geographically and ontogenetically. Specimens from the
Caribbean lowlands have blunt snouts in lateral view; those from the
Pacific lowlands have longer, more slender snouts that are pointed in
lateral view, and those from the Meseta Central are intermediate in
snout shape between the two lowland populations (Fig. 4). These
differences in shape of the snout are dependent on the nature of the
underlying cranial bones, principally the maxillaries and nasals. In
specimens from the Caribbean lowlands the nasals are long, wide, and
narrowly separated from the ethmoid; the anterior edge is just posterior
to the nostril. The maxillary flanges are nearly vertical. In specimens
from the Pacific lowlands the nasals are relatively shorter, narrower,
and rather widely separated from the ethmoid; the anterior edges of the
nasals do not extend so far forward as in specimens from the Caribbean
lowlands. The maxillary flanges slant medially. In these cranial
characters, specimens from the Meseta Central are intermediate between
the two lowland populations.

Superimposed on this geographic variation are ontogenetic changes, which
are most noticeable in males. In smaller, and presumably younger,
specimens the snouts are more pointed than in larger specimens;
consequently some small males from the Caribbean lowlands resemble
larger males from the Pacific lowlands, since the nasals and maxillaries
of the former are not fully ossified. In addition, in small breeding
males the ethmoid is only about one-half ossified, a large
frontoparietal foramen is present, the anterior arm of the squamosal
extends only about one-fourth the distance to the maxillary (two-thirds
the distance in larger specimens), and the tegmen tympani are short, as
compared with the long, thin elements in larger specimens.

    [Illustration: FIG. 4. Variation in the shape of the snout in
       _Smilisca sordida_; left column females, right column males;
       all from Costa Rica: (A) Camp Seattle, Rincón de Osa, Puntarenas
       Prov. (UMMZ 123684); (B) Quebrada Agua Buena, 3 km. SW Rincón de
       Osa, Puntarenas Prov. (USC 7236); (C) Río Oro, 28.5 km. NW Villa
       Neily, Puntarenas Prov. (KU 91742); (D) Río Jorco, near
       Desamparados, San José Prov. (KU 91765); (E-F) Bambú, Limón Prov.
       (USC 7183). ×3.]

The dorsal ground-color of _Smilisca sordida_ is gray to pale tan or
reddish brown; the venter is white. The dorsum is variously marked with
dark gray, dark brown, reddish brown, or olive-green spots or blotches
(Pl. 7C). A dark interorbital bar usually is present. The dorsal
markings on the body usually consist of a blotch, or two or more spots,
on the occiput, in the scapular region, and in the sacral region. In
many specimens, especially females, these markings are in the form of
broad transverse bars. A female (USC 7164) from Las Cañas, Guanacaste,
Costa Rica, has a tan dorsum with many black flecks and round brown
spots bordered by darker brown. One female (KU 91763) from the Río
Jorco, San José, Costa Rica, has a unicolor tan dorsum. Some individuals
have scattered, small white spots on the dorsum; these are most evident
in a male (USC 7153) from La Fortuna, Alajuela. White labial stripes and
anal stripes are absent in all specimens.

The limbs are marked by dark brown transverse bars; these are indistinct
in some specimens from the Meseta Central and Caribbean lowlands,
whereas they are distinct in all specimens from the Pacific lowlands.
Specimens from the Caribbean lowlands have two to six bars on each
shank, whereas specimens from the Pacific slopes have four to six bars
on each shank, and specimens from the Meseta Central have as many as
eight bars on each shank. A narrow, sometimes broken white line is
present on the ventrolateral edge of the forearm. The webbing on the
hand is tan or pale gray, and the ventral surfaces of the tarsi and the
webbing on the feet are dark gray or brown. Breeding males have dark
brown nuptial excrescences on the prepollex.

The flanks and posterior surfaces of the thighs usually are marked by
bluish white and creamy tan flecks, respectively, but vary considerably.
In specimens from the Caribbean lowlands a small amount of flecking is
present in the inguinal region, and on the posterior surfaces of the
thighs flecks are few or absent. In specimens from the Meseta Central,
numerous large flecks or small, round spots (pale bluish white in life)
are on the posterior half of the flanks; small flecks are on the
posterior surfaces of the thighs. Specimens from the Pacific slopes and
lowlands of southern Costa Rica (Puntarenas and San José Provinces) have
bold mottling of black and bluish white on the flanks and many bluish
white flecks on the posterior surfaces of the thighs. The flanks are
reticulated from the axilla to the groin in two females (UMMZ 123684 and
USC 7236) from Rincón, Peninsula de Osa. In specimens from the Pacific
slopes of Guanacaste in northwestern Costa Rica, flecks are present in
the inguinal region; indistinct flecks are on the posterior surfaces of
the thighs.

The throat is immaculate in specimens from the Caribbean lowlands in
Limón Province; the throats are dusky laterally in most other specimens
except some from the Meseta Central, in which the throats are heavily
flecked with black. This variation occurs in males and females.

The color and pattern in life are highly variable. A composite
description of living individuals (now KU 91718-41) from 6 kilometers
east of Golfito, Puntarenas, Costa Rica, illustrates the variability:
"Dorsum pale olive-green, fading to tan posteriorly, or tan all over
with dark olive-green or dark brown spots on back and bars on limbs.
Flanks dark brown with cream, greenish gray, or bluish gray mottling.
Posterior surfaces of thighs dark brown with pale blue, pale green, or
tan flecks. Iris creamy silver. Throats white with some brown flecks
peripherally." (Duellman, Field notes, February 15, 1965.) A male from
the Río Jorco, San José, Costa Rica, was dull olive-tan above with
olive-green marks; the flanks were brown with pale tan flecks, and the
posterior surfaces of the thighs were pale brown with cream-colored
flecks. Six females from the same locality were reddish brown above with
olive-brown or dark brown markings; one was uniform orange-tan, and
another was dull olive-green with darker markings.

The color of the iris in living frogs varies from creamy silver to
grayish yellow or bronze with a variable amount of black reticulation.

_Natural History._--_Smilisca sordida_ is not associated with any one
type of vegetation; instead it lives in the vicinity of rocky streams
having low gradients. Breeding takes place primarily in the dry season,
when the water in the streams is clear and at a low level. Through most
of the range of _S. sordida_ showers, or even short heavy rains, occur
in the dry season. After such rains the breeding activity is maximal.
Breeding congregations have been found from December through April, but
a few calling males and gravid females have been taken in June, July,
and August. In the rainy season non-breeding individuals are found
sitting on bushes near streams at night. Taylor (1952:843) found
specimens in bromeliads by day.

Males usually call from rocks or gravel bars in, or at the edge of,
streams. Some individuals perch in low bushes overhanging the streams,
and some sit in shallows in the streams. Clasping pairs have been found
on the banks of streams and in shallow water in streams.

The breeding call consists of one to six moderately short, rather
high-pitched notes (duration 0.18 to 0.45 seconds) repeated at intervals
of 12 seconds to several minutes. Each note is a vibrant rattle having
78 to 135 pulses per second and major frequences of about 1200 to 2600
cycles per second (Pl. 11C).

The tadpoles live in shallow parts of the streams, where they cling to
the surfaces of small rocks and hide beneath leaves and rocks. A
complete developmental series of tadpoles is not available; measurements
of those stages examined are summarized in Table 12.

A typical tadpole in stage 36 of development (KU 68475 from 15 km. WSW
of San Isidro el General, Costa Rica) has a body length of 11.7 mm.,
tail length of 22.8 mm., and a total length of 34.5 mm.; body about
three-fourths as deep as wide; snout broadly rounded in dorsal view,
sloping and rounded in lateral view; eyes widely separated, directed
dorsolaterally; nostril slightly closer to eye than to tip of snout;
mouth ventral; spiracle sinistral, about two-thirds distance from snout
to posterior end of body and slightly below midline; anal tube dextral;
caudal musculature heavy, straight; dorsal fin not extending onto body;
fins deepest at about mid-length of tail; there depth of caudal
musculature equal to depth of dorsal fin and half again as deep as
ventral fin; musculature extending nearly to tip of tail; body reddish
brown above and pale grayish brown with white flecks below; caudal
musculature pale tan with brown flecks; a series of reddish brown dashes
at base of caudal fin separated from others in series and from dashes on
other side by creamy white; fins transparent with reddish brown flecks
on posterior one-half of ventral fin and on all of dorsal fin (Fig.
14C). Mouth bordered by two rows of short, pointed papillae; lateral
fold present; tooth-rows 2/3; upper rows equal in length; second upper
row narrowly interrupted medially; three lower rows complete, nearly as
long as upper rows, deeply indented medially; upper beak robust, inner
surface not forming continuous arch with short lateral processes; lower
beak deep, V-shaped; both beaks bearing short serrations (Fig. 15F).

Little variation occurs in structure. In some specimens the second upper
tooth-row is complete; no individuals were found to have the row broadly
interrupted medially.

The series of dark dashes on the dorsal edge of the caudal musculature
is diagnostic of all stages studied. In life, tadpoles from 15 and 20
kilometers west-southwest of San Isidro el General, Costa Rica, had a
tan body, often with an olive-tan tinge; the caudal musculature was
tan; the flecks and dashes were dull red or reddish brown. Tadpoles from
6 kilometers east of Golfito, Costa Rica, had bodies with olive-green
flecks. The caudal musculature was brown with bluish green flecks; the
fins were transparent with reddish brown flecks. The belly was a silvery
golden color. Tadpoles from Bajos de Jorco, Costa Rica, had brown bodies
with bluish green flecks; the tail and fins had reddish brown flecks and
dashes. The iris was a bronze color in specimens from all three
localities, as well as in the young mentioned in the following
paragraph.

Nine recently metamorphosed young were found on vegetation at the edges
of streams in April. These specimens have snout-vent lengths of 13.1 to
15.7 mm. (14.9 mm.) and in life were pale greenish tan or olive-tan
above and white below. The hands, feet, and thighs were pale yellowish
tan.

_Remarks._--The foregoing synonymies indicate that confusion has existed
in the application of various names, to this species, as well as in use
of the names _sordida_ and _gabbi_ to include the species that we
describe and name _Smilisca sila_. Correct allocation of the names
involved was possible only after studying and comparing the type
specimens, for the descriptions given by the various authors are not
sufficiently explicit to determine the nature of many essential
features.

The presence of a rounded snout and a long white throat in males
distinguishes _S. sordida_ from _S. sila_, which has a high truncate
snout and short dark throat in males. The two syntypes of _Hyla sordida_
Peters, 1863, (ZMB 3141) are males having snout-vent lengths of 36.9 and
37.0 mm. The two syntypes of _Hyla gabbi_ Cope, 1876 (USNM 30658-9), are
females having snout-vent lengths of 52.8 and 53.7 mm., respectively.
Also included in the collections made by Gabb is eastern Costa Rica are
two males (USNM 30685-6), which Cope (1876) named and described as _Hyla
nigripes_. These specimens are soft and faded, but are recognizable as
the same as _Hyla sordida_ Peters; the syntypes of _Hyla nigripes_ have
snout-vent lengths of 37.6 and 37.7 mm. We have examined one of the
syntypes of _Hyla salvini_ Boulenger, 1882 (BMNH 1947.2.24.13), a female
having a snout-vent length of 54.6 mm. We are convinced that all of
these type specimens are representatives of one species, the earliest
name for which is _Hyla sordida_ Peters, 1863. The type localities for
three of the named species are in Costa Rica--_H. gabbi_ from Sipurio on
the Caribbean lowlands, _H. nigripes_ from the Caribbean slopes of Pico
Blanco, and _H. salvini_ from Cartago on the Meseta Central. The type
locality of _H. sordida_ was given as "Veraguas" by Peters (1863). At
that time Veraguas was often considered to be most of western Panamá.
Though we have not seen Panamanian specimens other than the types of _S.
sordida_ and one specimen from the Pacific lowlands of western Panamá,
the species probably occurs on the Caribbean slopes of western Panamá.
The species has been taken on the Caribbean lowlands of Costa Rica
within a few kilometers of Panamá; collecting on the Caribbean slopes in
the provinces of Bocas del Toro and Veraguas should reveal the presence
of _Smilisca sordida_ there.

_Distribution._--_Smilisca sordida_ is found along the Pacific slopes
and lowlands from Guanacaste, Costa Rica, southeastward to extreme
western Panamá, to elevations of about 1200 meters on the Meseta Central
in Costa Rica, and on the Caribbean slopes and lowlands of Costa Rica
and probably adjacent Panamá (Fig. 5). One specimen purportedly comes
from "Río Grande, Nicaragua."

    [Illustration: FIG. 5. Map showing locality records for
      _Smilisca sordida_.]

_Specimens examined._--412, as follows: NICARAGUA: "Río Grande"
(? Depto. Zelaya), MCZ 2634.

COSTA RICA: =Alajuela=: Between Atena and Salto de San Mateo, USC 6185;
8 km. N Ciudad Quesada, USC 7155 (4); La Fortuna, USC 7153 (20); 3 km. E
La Fortuna, USC 7150; San Carlos, USNM 29969; Sarchi, KU 32990-9,
36792-3.

=Cartago=: Cartago, BMNH 1947.2.24.13; headwaters of Río Pacuare, USC
119; Instituto Interamericano de Ciéncias Agricolas, Turrialba, KU
37012, USC 420, 437; Río Reventazón, Turrialba, MCZ 29268: 10 km. N Río
Reventazón bridge, USC 7073; 5 km. SW Río Reventazón bridge on
Paraiso-Orosi road, USC 669; Turrialba, UMMZ 118405, USC 455, USNM
29936-9.

=Heredia=: Puerto Viejo, KU 36791.

=Guanacaste=: Las Cañas, USC 7164; Santa Cecilia, MCZ 7924-5; Tilarán,
USC 7161 (5).

=Limón=: Bambú, USC 7171 (2), 7183 (13); La Lola, USC 820 (6), 6083-94,
8064, 8071; Pandora, USC 7188 (7), 7189, 7190 (3), 7191 (5); Pico
Blanco, USNM 30685-6; Río Larí, 14-16 km. SW Amubre, USC 7179, 7180
(10); Sipurio, USNM 30658-9; Suretka, KU 36764, 36765 (skeleton),
36766-78.

=Puntarenas=: 6 km. N Dominical, KU 91749-50, 91811 (young), 91812
(tadpoles); Esparta, MCZ 8028; 6 km. E Golfito, KU 91718-41, 91809
(young), 91810 (tadpoles), 91816-9 (skeletons), USC 7103 (23); Quebrada
Agua Buena, 3 km. SW Rincón de Osa, USC 7236 (6); Quebrada Boruca, 22
km. E Palmar Norte, KU 64264; Rincón de Osa, Camp Seattle, UMMZ
123680-5, S-2792 (skeleton), USC 705 (5), 6023, 7254; Río Barranca, USC
7119 (2); Río Ceiba, 6 km. NW Buenos Aires, KU 91747-8, USC 7112 (7);
Río Ciruelitas, 16 km. NW Esparta, USC 7121 (3); Río Claro, 14.2 km. NW
Villa Neily, USC 7110 (4); Río Ferruviosa, 7 km. S Rincón de Osa, USC
7235 (4); Río Lagarto at Pan-American Hwy. (Guanacaste Border), USC 7122
(4); Río La Vieja, 30 km. E Palmar Norte, KU 87684 (tadpoles), 91743-6,
USC 7083 (2); Río Oro, 28.5 km. NW Villa Neily, KU 91742; Río Volcán, 10
km. W Buenos Aires, USC 7113; Río Zapote, 7 km. E Palmar, USC 7100 (4);
3-5 km. W Palmar, USC 7101 (18); 7 km. SE Palmar, KU 64261-3; 1.2 km. NW
Villa Neily, USC 8032; 3 km. NW Villa Neily, USC 7109 (20); 5 km. NW
Villa Neily, USC 6176, 8035.

=San José=: Bajos de Jorco, KU 91813 (tadpoles); Escazú, KU 34863,
34869-75, USC 813; between Monrovia and La Hondura, ± 0.5 km. N Santa
Rosa, USC 302 (2); Paso Ancho, Río Jorco, UMMZ 122649 (6), USC 530 (3);
Río Jorco, near Desamparados, KU 91757-65, 91796-7, 91820-3 (skeletons),
USC 228, 513, 7117 (7); Río Peje, 10 km. SSE San Isidro el General, USC
7115 (3); Río Tiriví, MCZ 7972; San Isidro el General, CNHM 101096, KU
28201, 32989, UMMZ 72024; 15 km. WSW San Isidro el General, KU 64245-56,
68473 (tadpoles), 68474 (young), 68475 (tadpoles), 86516, 91754-6,
91793-5, USC 7097 (6); 17.1 km. WSW San Isidro el General, USC 6047; 18
km. WSW San Isidro el General, USC 689; 20 km. WSW San Isidro el
General, KU 64257-9, 64260 (skeleton), 68468 (young), 68469 (tadpoles),
68470 (young), 68471-2 (tadpoles), 68476 (young), 68633-4 (skeletons),
91751-3; San José, AMNH 7501-4, USC 298; Santa Rosa, Río Virilla, USC
7145.

PANAMA: =Chiriquí=: Río Jacu, 5.8 km. ESE Paso Canoas, KU 91905.
"Veraguas," ZMB 3141 (2).



ANALYSIS OF MORPHOLOGICAL CHARACTERS


Osteology

In attempting to assay the taxonomic significance of skeletal
differences we are faced with a dearth of data on the skeletons of frogs
in general and hylids in particular. Recent reviews by Brattstrom (1957)
and Hecht (1962, 1963) have been concerned with general salientian
classification and phylogeny, principally at the family level. Savage
and Carvalho (1953), Griffiths (1959), and Baldauf (1959) used
osteological characters in determining the taxonomic status of the
families Pseudidae, Brachycephalidae, and Bufonidae, respectively.
Carvalho (1954) presented osteological evidence for the generic
separation of New World microhylids. Zweifel (1956) and Tihen (1962)
used osteological characters at the levels of the species-group and
species in their respective studies on _Scaphiopus_ and _Bufo_. Little
has been recorded about the skeletons of the hylids. Goin (1961)
mentioned dentigerous elements and cranial co-ossification in his
synopsis of the genera of hylids. Copland (1957) in his review of
the _Hyla_ of Australia, Funkhouser (1957) in her revision of
_Phyllomedusa_, and Zweifel (1958) in his review of _Nyctimystes_ did
not consider skeletal characters.

Some osteological studies on hylids have yielded worthwhile information.
Mittleman and List (1953) used osteological characters in defining the
genus _Limnaoedus_: Starrett (1960) used cranial characters in
combination with jaw musculature in defining the genus _Smilisca_, and
Duellman (1964) used cranial characters in delimiting the _Hyla
bistincta_ group. Brief descriptions of cranial structure were given for
_Phrynohyas_ (Duellman, 1956) and _Ptychohyla_ (Duellman, 1963a);
specific and sexual differences in the skulls of _Hyla chaneque_ and
_Hyla taeniopus_ were pointed out by Duellman (1965). Stokely and List
(1954) described early cranial development in the hylid _Pseudacris
triseriata triseriata_.

Because our knowledge of the skeleton in hylids is so incomplete, we are
not attempting to place _Smilisca_ in the general scheme of hylid
phylogeny on the basis of skeletal characters. Instead, our purposes are
to describe the skeleton and its ontogenetic development in one member
of the genus (_S. baudini_), and to make comparisons that show taxonomic
differences in osteological characters among species of _Smilisca_.

The study of 68 dried skeletons and 25 cleared and stained preparations,
including an ontogenetic series of _S. baudini_, has resulted in an
understanding of the progressive development of skeletal elements and a
knowledge of interspecific and intraspecific variation in these
elements. Furthermore, investigations of the osteology have provided
correlations between some cranial characters and certain aspects of
external morphology.

_Descriptive Osteology of Smilisca baudini_

The following description is based primarily on an adult female (KU
68184):

_Skull._--The skull is large, solid, and broader than long; the greatest
width is between the sutures of quadratojugal and maxillary on either
side of the skull (Pls. 2-3). The maxillaries bear well-developed dorsal
flanges, curve gently, join the moderately convex premaxillaries
anteriorly and form a slightly truncate snout. The combined premaxillary
width is about one-fourth the width of the skull. The premaxillaries are
separated medially, and laterally from the maxillaries by sutures. Each
premaxillary bears a dorsomedial alary process, which is anteriorly
convex and four times as high as the depth of the lateral wing of
premaxillary; each premaxillary also has a ventromedial palatine process
that projects dorsally from the lingual edge of the premaxillary. The
septomaxillaries are closely associated dorsally with the premaxillaries
immediately lateral to the prenasal processes.

The nasals are large, widest anteriorly and narrowing posteriorly,
parallel to maxillaries, and not separated from the ethmoid by
cartilage. The nasals bear long, delicate maxillary processes extending
nearly to the maxillaries. Anteriorly, the nasals are widely separated
by the partially ossified internasal septum, which is in contact with
the premaxillaries between the prenasal processes; the anterior points
of the nasals lie approximately one-half the distance between the
anterior ends of the ethmoid and the premaxillaries. The ethmoid is
large and completely ossified; the margins are smooth. The trunate
anterior edge lies between the nasals and is in contact with the
internasal septum. The frontoparietals are large, smooth-margined, and
bear large supraorbital flanges curving posterolaterally at the rear of
the orbit. A small, oval foramen involves the posterior part of the
ethmoid and anterior portion of frontoparietals; continued ossification
in older specimens fills in the foramen, thereby resulting in a solidly
roofed cranium. The auditory regions are relatively massive and bear
narrow tegmen tympani; the distal ends of the tegmen tympani are medial
to the lateral edge of the pterygoids in dorsal view. The squamosals are
large; the long anterior arm is separated from the maxillary by a
suture. The delicate, spindle-shaped columellae lie ventral to the
tegmen tympani and squamosals, are spatulate distally, and have a broad
basal attachment to the auditory region.

The vomers are moderately large and are in contact anteriorly with the
premaxillaries and posteriorly with the ethmoid. Each vomer has two wide
serrated flanges laterally. The tooth-bearing parts of the vomers are
widely separated and at a slight angle to one another; the vomers
terminate medially in two pointed processes on the ethmoid. The
palatines are edentate, but bear strong ridges throughout their lengths.
They are broadly in contact with the maxillary, are narrow medially, and
are attached by pointed processes to the medial part of the ethmoid.
The pterygoids are large, attached to the maxillaries immediately
anterior and medial to the squamosal-maxillary connection, bear
well-developed pedicles, which are broadly attached to the proötic, and
a wide wing is in contact posteriorly with the distal two-thirds of the
quadrate.

The angular makes up most of the lower jaw, bears a broad articular
surface posteriorly, and has a small coronoid process on the lingual
edge; anteriorly the angular is separated from the dentary and
mentomecklian by Meckel's cartilage. The dentary lies external to the
angular and extends from the mentomecklian to approximately the
mid-length of the angular. The mentomecklians are ossified, but
separated by cartilage medially.

_Hyoid._--The hyoid plate is curved, thin, and mostly cartilaginous, but
calcined posteriorly (Fig. 6). The anterior cornua are slender,
cartilaginous, and curve anteromedially from the hyoid plate and thence
laterally and posteriorly, to attach to the posterior surface of the
proötics. The lateral cornua are broad, flat, cartilaginous lateral
extensions from the bases of the anterior cornua. The posterior cornua
are bony, except distally.

    [Illustration: FIG. 6. Ventral view of hyoid apparatus of an adult
       male _Smilisca baudini_ showing areas of muscle attachment: _Gen.
       L._, attachment of geniohyoideus lateralis; _Gen. M._, attachment
      of geniohyoideus medialis; _Hyo._, attachment of hyoglossus; _Omo._,
       attachment of omohyoideus; _Pet._, petrohyoideus; _St._, attachment
       of sternohyoideus. KU 64220, ×5.]

_Vertebral Column._--The atlas lacks transverse processes and a neural
crest, whereas transverse processes are present on the other seven
presacral vertebrae, and knoblike neural crests are present on the
second, third, and fourth vertebrae; a faint neural ridge is visible on
the fifth vertebra. The transverse processes are directed laterally on
the second and sixth vertebrae, ventrolaterally on the third,
posterolaterally on the fourth and fifth, and anterolaterally on the
seventh and eighth. The processes are slightly expanded on the fourth,
and more so on the fifth, vertebra. The sacral diapophyses are expanded
and have a border of calcified cartilage laterally. There are two sacral
condyles. The slender coccyx has a thin dorsal ridge on the anterior
three-fourths of its length.

_Pectoral Girdle._--The omosternum is large, ovoid, and cartilaginous;
the sternum is a thin cartilaginous sheet deeply notched posteriorly and
is not differentiated into episternal and xiphisternal elements. The
coracoids are robust, twice as stout as the clavicles. The epicoracoidal
cartilages overlap in the usual arciferal manner, except that they are
fused anteriorly between the slender clavicles. The clavicles are
strongly arched. The clavicle, coracoid, and scapula on each side form a
bony articulation at the glenoid fossa. A bifurcation of the ventral end
of the scapula results in a large glenoid foramen. The scapula is flat
and expanded dorsally; the suprascapula is broad, flat, and calcified in
large adults. In young specimens no distinct ossification of the
cleithrum or ossification of endochondral centers are evident.

_Arm and Hand._--The humerus is equally well-developed in both sexes and
has a prominent lateral crest. The radius and ulna are completely fused.
A bony prepollex is present in both sexes. The metacarpals are about
equal in length. The phalangeal formula is 2-2-3-3; the terminal
phalanges are claw-shaped.

_Pelvic Girdle._--The ilia are long, slender, and slightly curved. A
thin ridge projects laterally from the dorsal edge of the posterior
one-half of each ilium. The ilial prominence is large and knoblike when
viewed from above. The anterior edge of the ilial prominence is at the
level of the anterior edge of the acetabular border. The dorsal
acetabular expansion is small. The pubis is slender, and the ischium is
elevated and robust.

_Leg and Foot._--The slightly curved femur has a distinct crest
proximally on the posterior surface. The nearly straight tibio-fibula is
slightly longer than the femur. The tibial and fibial elements are
completely fused but have a distinct cleft between them. A small foramen
exists at the mid-length of the tibio-fibula. The fibulare (calcaneum)
is much more robust than the tibiale (astragalus). The prehallux is
large and flat. The metatarsals of the third, fourth, and fifth digits
are equal in length; the metatarsal of the second is somewhat shorter,
and that of the first is much shorter. The phalangeal formula is
2-2-3-4-3; the terminal phalanges are claw-shaped.

_Developmental Cranial Morphology of Smilisca baudini_

The following description of development of the skull of _Smilisca
baudini_ is based on the examination of 12 cleared and stained
specimens. In table 3 the cranial bones are listed in the left hand
column in the approximate order of their appearance in the young frogs.
Across the top of the table selected specimens designated by
developmental stage or snout-vent length are listed. It should be noted
that although each individual, from left to right, has an increasing
number of ossified bones, the correlation with increasing size is
imperfect; the precise ages of the individuals are unknown.

The first bones to appear are the septomaxillaries, frontoparietals,
part of the exoccipital, and the parasphenoid in developmental stage 40.
The frontoparietals are represented by two slender ossifications
dorsomedial to the orbits; the septomaxillaries are present as small
ossifications anterior to the nasal capsules (Pl. 1A). The parasphenoid
is present as a faint median ossification, and the exoccipital shows
some ossification.


  Table 3.--The Order of Occurrence of Cranial Ossifications in the
     Skull of Smilisca baudini. Where Numbers Are Divided by a Slash
     Mark, the Left and Right Symbols Correspond to the Left and
     Right Sides of the Skull, Respectively.

  =====================+=====+=====+=====+=====+=====+=====+====
  Bone                 |Stage|Stage|12.6 |13.9 |32.0 |27.0 |20.1
                       |  40 |  44 | mm. | mm. | mm. | mm. | mm.
  ---------------------+-----+-----+-----+-----+-----+-----+----
  Frontoparietal       |  X  |  X  |  X  |  X  |  X  |  X  |  X
  Parasphenoid         |  X  |  X  |  X  |  X  |  X  |  X  |  X
  Septomaxillaries     |  X  |  X  |  X  |  X  |  X  |  X  |  X
  Exoccipitals         |  X  |  X  |  X  |  X  |  X  |  X  |  X
  Squamosals           | --  |  X  |  X  |  X  |  X  |  X  |  X
  Premaxillaries       | --  |  X  |  X  |  X  |  X  |  X  |  X
  Maxillaries          | --  |  X  |  X  |  X  |  X  |  X  |  X
  Nasals               | --  | --  |  X  |  X  |  X  |  X  |  X
  Pterygoids           | --  | --  |  X  |  X  |  X  |  X  |  X
  Vomers               | --  | --  | --  |  X  |  X  |  X  |  X
  Palatines            | --  | --  | --  |  X  |  X  |  X  |  X
  Quadratojugals       | --  | --  | --  |  X  |  X  |  X  |  X
  Ethmoid              | --  | --  | --  | --  |  X  |  X  |  X
  Columellas           | --  | --  | --  | --  |  X  |  X  |  X
  Supraorbital Flanges | --  | --  | --  | --  | --  |  X  |  X
  Proötics             | --  | --  | --  | --  | --  | --  |  X
  Vomerine Teeth       | --  | --  | 1/1 | 4/3 | 5/5 | 3/3 | 5/4
  Maxillary Teeth      | --  | 0/7 | 3/5 | 6/5 |30/31|30/26|37/36
  Premaxillary Teeth   | --  | 2/4 | 3/3 | 5/5 | 7/6 | 8/6 | 8/7
  ---------------------+-----+-----+-----+-----+-----+-----+----


The dentigerous bones are among the most rapidly developed, although not
the first to appear. They are present in developmental stage 44 before
metamorphosis is completed. The maxillaries bear a few teeth anteriorly
and are ossified posteriorly to a point one-third of the distance from
the anterior to the posterior edge of the orbit. Ossification lengthens
the posterior termini of the maxillaries to the posterior edge of the
orbit. In front of the anterior margin of the orbit, bone is
proliferated dorsal to the main axes of the maxillaries and forms
moderate dorsal maxillary flanges. The premaxillaries appear
simultaneously with the maxillaries. Initially they are widely separated
medially from each other, and laterally from the developing maxillaries;
each bears two or three teeth, large dorsally blunt alary processes, and
small palatine processes. The median and lateral edges of the prenasal
processes lengthen heterochronously, causing the median edges to be
longest and to lie slightly dorsal to the level of the septomaxillaries.
After the maxillaries and premaxillaries develop, the vomers appear as
small horizontal ossifications anterior to the parasphenoid.
Ossification begins in the lateral flanges, then in the prevomerine
processes, and lastly in the posterior dentigerous parts of the bones;
the prevomerine processes are the last parts of the vomers to ossify
completely.

Initially the frontoparietals are present as thin rods of ossification
dorsomedial to the orbits; the frontoparietals extend from the anterior
to the posterior end of the orbit by developmental stage 44. The
anterior ends of the bones remain thin and pointed; ossification
progresses medially from the midpoint of the length of the orbit and
posteriorly to the level of the exoccipital; a median center of
ossification joins the frontoparietals posteriorly, thereby forming the
posterior border of the frontoparietal fontanelle. The supraorbital
flanges of the frontoparietals do not appear until all other cranial
bones are ossified, or nearly so. The most rapid ossification begins
laterally at the posterior edge of the orbit and decreases anteriorly
over the posterior half of the orbit. This differential rate of
proliferation of bone results in the pattern of development of the
supraorbital flanges shown in figure 7. The nasals appear as
thin slivers of bone half way between the anterior ends of the
frontoparietals and the end of the snout. As ossification proceeds the
nasals assume a triangular shape in dorsal view. The anterior ends are
pointed; the lateral margins are parallel to the maxillaries. The
posteromedial points do not reach the lateral margins of the ethmoid,
and the maxillary processes extend about three-fourths the distance from
the bodies of the nasals to the maxillaries. Following the union of the
frontoparietals posteriorly, the nasals widen anteriorly and are
narrower at the midpoints of their long axes than anteriorly or
posteriorly. With further ossification the maxillary processes extend to
the maxillaries and form complete bony anterior margins to the orbits;
the mid-parts of the nasals widen (Pl. 1B).

    [Illustration: FIG. 7. Developmental sequence of the frontoparietal
       fontanelle and associated bony elements in _Smilisca baudird_:
       (A) KU 60026, ×5; (B) KU 85438, ×4; (C) KU 26328, ×3;
       (D) KU 68184, ×2.3.]

The parasphenoid is the first of the palatal bones to appear. At
metamorphosis the bone is well developed; the anterior tip is situated
just in front of the anterior edge of the orbit, and posteriorly the
lateral processes extend laterally beyond the ossified parts of the
auditory region. The pterygoids do not appear until metamorphosis, when
ossification is evident in only the mid-parts of the posterolateral
arms. Ossification follows in the mid-parts of the anterolateral arms
and occurs last in the pterygoid pedicles. The palatines do not appear
until all three arms of the pterygoids are at least partly ossified.
Ossification proceeds rapidly from the maxillaries medially to the
unossified ethmoid, which is the last of the cranial bones to appear.
Initially it is extremely shallow; dorsally it is widely separated from
the nasals, and ventrally the posterior margin meets the anterior point
of the parasphenoid. In dorsal view, ossification proceeds anteriorly
between the nasals and posteriorly, ventral to the frontoparietals;
ventrally, ossification proceeds posteriorly dorsal to the parasphenoid.

The ventral arms of the squamosal and the supraoccipital region of the
exoccipital are the first occipital bones to appear. Ossification
follows in the regions of the semicircular canals and occipital
condyles. The dorsal end of the ventral arm of the squamosal and the
posterior arm of the squamosal ossify as a unit at the same time the
quadratojugal appears. Shortly thereafter the anterior arm of the
squamosal ossifies, the distal part of the columella appears, and the
anterior and lateral parts of the auditory region ossify.

The angular and dentary of the lower jaw appear concurrently with the
dentigerous bones. Initially, the angular is short and broad; the
articular surface is absent, and the anterior end is slightly overlapped
by the dentary. The mentomecklians do not ossify until approximately
the same time that the quadratojugal appears in the upper jaw.

_Comparative Osteology_

The genus _Smilisca_ is characterized by the following combination of
cranial osteological characters: (1) A large amount of bone is involved
in the skull and a minimal amount of cartilage and/or secondarily
ossified cartilage; co-ossification is absent. (2) The skulls are
uniformly broad with angular lateral margins, and truncate anteriorly.
(3) An internasal septum and quadratojugals are present. (4) A
well-developed squamosal minimally extends one-fourth the distance from
the dorsal end of the quadrate to the maxillary, and maximally is
separated from the maxillary by a suture. (5) The ethmoid is large; the
distance between the anterior end of the ethmoid and the anterior edge
of the premaxillary varies between 15 and 20 per cent of the total
length of the skull.

On the basis of cranial osteology two species-groups can be recognized
within the genus _Smilisca_. The _sordida_ group, comprising _S.
sordida_ and _puma_, is characterized by a broad skull in which the
lateral margins of the maxillaries are relatively straight anterior to
the orbit. The moderate-sized nasals are rounded anteriorly, and bear
relatively short, sometimes blunt, maxillary processes. The long axes of
the nasals are not parallel to the maxillaries. The ethmoid is
proportionately small in the _sordida_ group. The bony part of the
ethmoid terminates near the anterior edge of the orbits and does not
extend anteriorly between the nasals; the entire anterior margin of the
ethmoid is separated from the nasals by cartilage. The squamosals are
generally small. They are narrow in dorsal view, and minimally extend
one-fourth the distance from the dorsal end of the quadrate to the
maxillary, and maximally, two-thirds the distance. The tegmen tympani
are relatively small (Fig. 8).

    [Illustration: FIG. 8. Dorsal views of the skulls of the species of
       _Smilisca_: (A) _S. baudini_ (KU 68184); (B) _S. puma_ (KU 68636);
       (C) _S. phaeota_ (KU 41090); (D) _S. sila_ (KU 80625); (E) _S.
       cyanosticta_ (KU 55938), and (F) _S. sordida_ (KU 36765). ×1.5.]

In contrast to the tendency for reduction of cranial parts in the
_sordida_ group, the _baudini_ group, constituted by _S. cyanosticta_,
_phaeota_, and _baudini_, is characterized by more ossification of the
cranial elements. The skull is broad; the lateral margins are less
angular and are gently curved, rather than straight as in the _sordida_
group. The nasals tend to be larger with the long axes parallel to the
maxillary. Anteriorly the nasals are pointed, and posteriorly they bear
long, delicate palatine processes extending to the maxillary. The
ethmoid is fully ossified, extends anteriorly between the nasals, and
laterally is separated by a suture from the nasals if the latter are
fully ossified. The squamosals are large, and wide in dorsal view. They
minimally extend one-fourth the distance from the dorsal end of the
quadrate to the maxillary, and maximally are sutured to the maxillary.
The tegmen tympani are massive.

_Smilisca sila_ is intermediate between the two species-groups
described. The skull is broad; the lateral margins are gently curved,
and have a pronounced angularity just anterior to the palatines which
results in a broad, truncate snout. The nasals are moderate in size;
because of the anterior angularity of the lateral margins, the long axes
of the nasals lie parallel to the maxillary. The nasals are only
slightly pointed anteriorly, and posteriorly they bear short, blunt
palatine processes and medial processes in contact with the lateral
corners of the ethmoid. The ethmoid is fully ossified, but does not
extend anteriorly between the nasals. The squamosals are moderate in
size and extend one-fourth the distance from the dorsal end of the
quadrate to the maxillary. The tegmen tympani are relatively large, but
proportionately short.

The cranial characters utilized in the analysis of species groups
(general shape, nature of the nasals, ethmoid, squamosals, and tegmen
tympani), together with other characters, such as the relative height
and shape of the prenasal processes, the extent of the internasal
septum, and the nature of the vomers, frontoparietals, maxillaries and
pterygoids are useful in distinguishing the various species (Table 4,
Fig. 8), as well as in establishing relationships within the
species-groups.

Within the _sordida_ group, _S. sordida_ and _S. puma_ can be
distinguished by the following characters: The bony part of the ethmoid
terminates posterior to the anterior edge of the orbit and is thus
widely separated from the nasals by cartilage in _S. puma_. In _S.
sordida_ the bony part of the ethmoid always terminates at a level equal
to, or slightly in front of the anterior edge of the orbit; therefore,
less cartilage exists between the ethmoid and nasals in _S. sordida_
than in _S. puma_. The width of the premaxillary comprises about 30 per
cent of the width of the skull in _S. sordida_ and 20 per cent in _S.
puma_. The proportion of the length of the skull anterior to the bony
part of the ethmoid in _S. sordida_ is approximately 21 per cent, as
compared with about 29 per cent in _S. puma_. The prenasal processes are
convex in _S. sordida_ and straight in _S. puma_.

The marked ontogenetic variation in _S. sordida_ is considered in more
detail in the account of that species, but it is pertinent to the
present discussion to note that with respect to some features of the
skull some young breeding specimens of _S. sordida_ are intermediate in
appearance between large females of _S. sordida_ and adults of _S.
puma_. In some breeding males (usually the smaller individuals) of _S.
sordida_ the bony part of the ethmoid terminates at the anterior edge of
the orbit and is widely separated from the nasals by cartilage. In small
individuals _S. sordida_, especially in males, and in adults of _S.
puma_ the tegmen tympani are relatively short, whereas in adult females
of _S. sordida_ these elements are long and slender. In the smaller
specimens of _S. sordida_ and in _S. puma_ the squamosal is small; it
extends only about one-fourth of the distance to the maxillary in the
smaller _S. sordida_ and about one-half the distance in _S. puma_. The
more massive squamosal in large adult females of _S. sordida_ extends at
least two-thirds of the distance to the maxillary.


  Table 4.--Comparative Cranial Osteology of Smilisca.

  ===============+==============================+=======================
    Character    |         _S. baudini_         |   _S. cyanosticta_
  ---------------+------------------------------+-----------------------
                 |                              |
  Alary Processes| Four times as high as        | Three times as high
                 | lateral wing of premaxillary;| as lateral wing of
                 | anteriorly                   | premaxillary;
                 | convex.                      | anteriorly
                 |                              | convex.
                 |                              |
  Nasals         | Long, wide anteriorly,       | Long, widest
                 | narrowing posteriorly;       | posteriorly;
                 | attached to ethmoid.         | attached to
                 |                              | ethmoid.
                 |                              |
                 |                              |
                 |                              |
  Ethmoid        | Long; entirely ossified;     | Long, entirely
                 | smooth margins.              | ossified;
                 |                              | smooth margins.
                 |                              |
  Frontoparietal | Small, ovid fontanelle       | Large fontanelle, two
                 | present or absent;           | and one-half times as
                 | long, pointed postorbital    | long as wide; narrow
                 | processes curving            | supraorbital flanges
                 | along posterior              | with irregular margins.
                 | border of orbit.             |
                 |                              |
  Squamosal      | Large: anterior arm          | Large; anterior arm
                 | in contact with maxillary.   | in contact with
                 |                              | maxillary.
  ---------------+------------------------------+------------------------

  TABLE 4 (Continued)
  ===============+=============================+=========================
    Character    |        _S. phaeota_         |          _S. puma_
  ---------------+-----------------------------+-------------------------
                 |                             |
  Alary Processes| Two and one-half            | Two times as high as
                 | times as high as lateral    | lateral wing of
                 | wing of premaxillary;       | premaxillary;
                 | anteriorly convex.          | straight.
                 |                             |
                 |                             |
  Nasals         | Long, widest anteriorly     | Short, narrow, not
                 | and posteriorly,            | attached to ethmoid.
                 | bearing posteromedial       |
                 | process; not attached       |
                 | to ethmoid.                 |
                 |                             |
                 |                             |
  Ethmoid        | Long, entirely ossified;    | Short, about two-thirds
                 | smooth margins.             | ossified; irregular
                 |                             | margins.
                 |                             |
  Frontoparietal | Fontanelle absent;          | Keyhole-shaped fontanelle;
                 | large supraorbital          | smooth margins;
                 | flanges having              | flanges absent.
                 | straight edges and extending|
                 | posterolaterally.           |
                 |                             |
                 |                             |
  Squamosal      | Large; anterior arm         | Small; anterior arm
                 | extending 1/2-2/3 way       | extending 1/2 way to
                 | to maxillary.               | maxillary.
  ---------------+-----------------------------+--------------------------

  TABLE 4 (Continued)
  ===============+===========================+============================
    Character    |       _S. sila_           |     _S. sordida_
  ---------------+---------------------------+----------------------------
                 |                           |
  Alary Processes| One and one-half          | Two and one-half
                 | times as high as lateral  | times as high as lateral
                 | wing of premaxillary;     | wing of premaxillary;
                 | straight.                 | slightly convex
                 |                           | anteriorly.
                 |                           |
  Nasals         | Short, wide, bearing      | Moderately long narrowest
                 | small posteromedial       | anteriorly and
                 | processes; not attached   | posteriorly; not attached
                 | to ethmoid.               | to ethmoid.
                 |                           |
                 |                           |
                 |                           |
  Ethmoid        | Moderately long; entirely | Short; one-half to entirely
                 | ossified; smooth          | ossified; irregular
                 | margins.                  | margins.
                 |                           |
  Frontoparietal | Large, ovoid fontanelle;  | Large, elongate fontanelle;
                 | smooth margins;           | smooth margins;
                 | flanges absent.           | flanges absent.
                 |                           |
                 |                           |
                 |                           |
                 |                           |
  Squamosal      | Moderately large; anterior| Moderately small; anterior
                 | arm extending             | arm extending
                 | 1/4 way to maxillary.     | 1/4-2/3 way to maxillary.
  ---------------+---------------------------+----------------------------


Within the _baudini_ group, the skull of _S. cyanosticta_ is the most
generalized of the three species; the cranial characters are
intermediate between _S. phaeota_ and _S. baudini_. The lateral margins
of the skull in _S. cyanosticta_ are gently curved, and have an
angularity anterior to the palatine-maxillary suture; the anterior
margins are less angular in _S. phaeota_, which has a broader snout.
Posteriorly in _S. baudini_ the margins are slightly curved
medially, and the greatest width of the skull is between the
quadratojugal-maxillary sutures on either side of the skull. The
frontoparietals of _S. cyanosticta_ bear slightly irregular lateral
margins and a large fontanelle. There is a tendency for obliteration of
the fontanelle with increasing age in both _S. baudini_ and _S.
cyanosticta_; the lateral margins of the frontoparietals bear large
supraorbital flanges in both of these species. In _S. phaeota_ the
flanges are most prominent; they extend posterolaterally with straight
margins along two-thirds of the length of the orbit and terminate in
rather blunt points. The broad interorbital flanges result in a
relatively broad external interorbital distance. In _S. baudini_ the
flanges are curved posterolaterally around the orbit and terminate in
sharp, thin points. The tegmen tympani of all three species are massive.
In _S. cyanosticta_ the proötics slope posteriorly, whereas they slope
anteriorly in _S. baudini_ and _S. phaeota_.

The skulls of _S. cyanosticta_ and _S. baudini_ are alike in certain
respects. The squamosals of both species are large and connected to the
maxillary by a bony connection; the squamosals of _S. phaeota_ are
large, but extend only two-thirds of the distance from the dorsal end of
the quadrate to the maxillary. In _S. baudini_ and _S. cyanosticta_ the
nasals are separated throughout their lengths from the ethmoid, whereas
the nasals of _S. phaeota_ are separated from the ethmoid by cartilage.
The latter separation is due to an incomplete ossification of the nasals
in _S. phaeota_. The bony part of each nasal is constricted in the
middle of the long axis of the bone, and the nasals are widest
anteriorly; posteriorly each nasal bears a medial process, which is
narrowly separated from the lateral edge of the ethmoid.


  Table 5.--Variation in the Number of Teeth in the Species of
     Smilisca. (All Are Males; N = Number of Jaws, or Twice the Number
     of Individuals; Means Are Given in Parentheses After the Observed
     Ranges.)

  =================+====+==============+==============+===========
  Species          |  N | Maxillary    | Premaxillary | Vomerine
  -----------------+----+--------------+--------------+-----------
  _S. baudini_     | 20 | 49-65 (56.0) |  9-16 (13.6) | 5-9 (7.2)
  _S. cyanosticta_ |  8 | 50-64 (57.9) | 10-12 (10.8) | 4-11 (7.1)
  _S. phaeota_     | 20 | 50-68 (58.1) | 10-15 (12.1) | 5-9  (7.3)
  _S. puma_        |  6 | 60-67 (63.6) | 11-13 (12.0) | 4-7 (5.3)
  _S. sila_        |  8 | 48-60 (52.9) | 10-14 (11.3) | 5-7 (5.7)
  _S. sordida_     | 12 | 39-55 (44.2) |  7-11 (9.3)  | 4-6 (5.2)
  -----------------+----+--------------+--------------+-----------


The teeth of all species of _Smilisca_ are spatulate and bifid. The
numbers of maxillary, premaxillary, and vomerine teeth are summarized in
Table 5. Smaller and presumably younger specimens of all species of
_Smilisca_ have fewer teeth than do larger specimens of the same
species. This correlation between size and number of teeth does not
exist as an interspecific trend within the genus; for example, the
smallest species in the genus, _S. puma_, has the highest number of
maxillary teeth. In small specimens of a given species wide gaps are
present between the maxillary teeth posteriorly; in large specimens the
gaps are filled by teeth, beginning anteriorly and progressing
posteriorly, until the maxillary dentition is continuous.


Musculature

No extensive study of the muscular system was undertaken, but certain
muscles know to be of taxonomic importance were studied.

_Jaw Musculature._--Starrett (1960) pointed out the unique jaw
musculature in _Smilisca_. In this genus M. depressor mandibulae
consists of two parts, one arising from the dorsal fascia and one from
the posterior arm of the squamosal. Two muscles arise from the anterior
arm of the squamosal and insert on the lateral face of the mandible. Of
these muscles, M. adductor mandibulae posterior subexternus lies medial
to the mandibular branch of the trigeminal nerve; the other, M. adductor
mandibulae externus superficialis, lies lateral to the same nerve (Fig.
9). In most other hylids the latter muscle is absent. No significant
variation in the position of the muscles was noted in the various
species of _Smilisca_, though M. adductor mandibulae originate somewhat
more anteriorly in _S. baudini_ and _S. cyanosticta_ than in the other
members of the genus, all of which have a shorter anterior arm of the
squamosal that does not reach the maxillary. The two separate parts of
M. depressor mandibulae are not so widely separated in members of the
_sordida_ group as in the _baudini_ group.

    [Illustration: FIG. 9. Lateral view of the left jaw of _Smilisca
       baudini_; _A. M. E. S._, adductor mandibulae externus
       superficialis; _A. M. P. S._, adductor mandibulae posterior
       subexternus; _Col._, columella; _D. M._ depressor mandibulae;
       _M. B. T. N._, mandibular branch trigeminal nerve;
       _Sq._, squamosal. KU 64214, ×5.]

    [Illustration: FIG. 10. Ventral view of throat musculature in an
       adult male _Smilisca baudini_ (Superficial musculature on left,
       deep musculature on right); _A. C._ anterior cornua of hyoid;
       _Gen. L._, geniohyoideus lateralis; _Gen. M._, geniohyoideus
       medialis; _Hyo._, hyoglossus; _Omo._, omosternum; _Pet._,
       petrohyoideus; _S._, submentalis; _Sm._, submaxillaris;
       _St._, sternohyoideus; _V. S._, vocal sac. KU 64220, × 2.5.]

_Throat Musculature._--The frogs that comprise the genus _Smilisca_ are
characterized by paired subgular vocal sacs, essentially the same as
those in _Triprion_ (Duellman and Klaas, 1964). The following
description is based on _Smilisca baudini_ (Fig. 10).

M. submentalis lies in the anterior angle of the lower jaw, is thick,
and consists of transverse fibers extending between the dentaries. M.
submaxillaris is thin and arises from the whole of the inner surface of
the lower jaw, except for the anterior angle occupied by M. submentalis.
Anteriorly M. submaxillaris is broadly attached by fascia to M.
hyoglossus and M. geniohyoideus, which lie dorsal to M. submaxillaris.
Medially this attachment continues posteriorly for about one-half the
length of the hyoglossus. Posteriorly M. submaxillaris is folded and
attached to M. sternoradialis of the pectoral girdle. The vocal sacs are
formed by a pair of posterolateral evaginations of M. submaxillaris; a
broad connection between the pouches allows free passage of air between
the pouches.

The deeper throat musculature is essentially the same as that described
for _Phrynohyas spilomma_ by Duellman (1956), except for slight
differences in the place of attachment on the hyoid.


Skin

_Structure_

The skin of _Smilisca_ is typical of that of most hylids in organization
and structure. _Smilisca sila_ is distinguished from other members of
the genus by the presence of small wartlike protrusions and peculiar
white, pustular spots on the dorsum. The wartlike structures are
composed of three or four epidermal cells, which protrude from the
surface of the epidermis; the structures are covered by a slightly
thickened layer of keratin. The white pustules are slightly elevated
above the surrounding skin. Internally they consist of aggregations of
swollen, granular, pigment-cells (perhaps lipophores) lying between the
epidermis and the melanophores.

_Biochemical Variations_

Dried skins of all species of _Smilisca_ were sent to José M. Cei,
Instituto Nacional de Cuyo, Mendoza, Argentina, for biochemical
screening by means of the chromatographic techniques described by
Erspamer and Cei (1963). The species in the _baudini_ group have
detectable amounts of penta-hydroxi-trypatamine, whereas only a trace is
present in the other species. Furthermore, species in the _baudini_
group differ from _S. sila_ and the _sordida_ group in lacking, or
having only a trace of, tryptophan-containing polypeptides. These
superficial biochemical tests support the arrangement of species as
ascertained by conventional taxonomic characters.


External Morphological Characters

The features of external morphology that were studied in connection with
the taxonomy of the genus _Smilisca_ are discussed below.

_Size and Proportions_

The frogs of the genus _Smilisca___ are medium to large tree frogs. The
three species comprising the _baudini_ group (_S. baudini_,
_cyanosticta_, and _phaeota_) are notably larger than _S. puma_, _sila_,
and _sordida_ (Table 6). The largest specimen that we examined is a
female of _S. baudini_ having a snout-vent length of 90 mm. _Smilisca
puma_ is the smallest species; the largest male has a snout-vent length
of 38 mm. and the largest female, 46 mm.


  Table 6.--Comparison of Sizes and Certain Proportions of the Species
     of Smilisca. (Means in Parentheses Below Observed Ranges; Data for
     Males Only.)

  ================+====+===========+=============+===========+
                  |    | Snout-vent|Tibia length/| Tympanum/ |
      Species     | N  |   length  |  snout-vent |    eye    |
  ----------------+----+-----------+-------------+-----------+
                  |    |           |             |           |
  _S. baudini_    |140 | 47.3-75.9 | 42.1-53.6   | 56.1-94.4 |
                  |    |  (58.7)   |  (47.8)     |  (73.5)   |
                  |    |           |             |           |
  _S. cyanosticta_| 40 | 44.6-56.8 | 51.9-59.7   | 62.7-88.4 |
                  |    |   (50.7)  |  (56.0)     |  (71.4)   |
                  |    |           |             |           |
  _S. phaeota_    | 50 | 40.8-65.5 | 50.9-60.2   | 62.7-85.5 |
                  |    |  (53.9)   |  (55.5)     |  (76.6)   |
                  |    |           |             |           |
  _S. puma_       | 20 | 31.9-38.1 | 48.2-53.1   | 52.1-72.2 |
                  |    |  (34.7)   |  (51.3)     |  (64.9)   |
                  |    |           |             |           |
  _S. sila_       | 33 | 31.6-44.8 | 49.7-58.1   | 47.6-58.3 |
                  |    |  (37.7)   |  (54.8)     |  (53.2)   |
                  |    |           |             |           |
  _S. sordida_    | 55 | 31.9-44.6 | 50.5-57.1   | 46.5-57.1 |
                  |    |  (37.9)   |  (53.4)     |  (49.1)   |
  ----------------+----+-----------+-------------+-----------+


No outstanding differences in proportions exist between species,
although certain proportions are sufficiently different in some species
to warrant mention. _Smilisca baudini_ is a more squat and stocky frog
than other members of the genus; this is reflected in the somewhat
shorter hind legs (Table 6). The size of the tympanum relative to that
of the eye is highly variable within samples of a given species. Even
so, noticeable differences in the tympanum/eye ratio are apparent.
Members of the _baudini_ group have the largest tympani, whereas _S.
sila_ and _sordida_ have the smallest, and _S. puma_ is intermediate
(Table 6).

_Shape of Snout_

Although all members of the genus have rather truncate snouts, subtle
differences exist among the species (Pl. 12). _Smilisca sila_ has the
shortest snout; that of _S. baudini_ is only slightly longer. The snouts
of _S. cyanosticta_ and _puma_ are nearly square in lateral profile,
whereas those of _S. phaeota_ and _sordida_ are slightly inclined. The
shape of the snout is relatively uniform within each species and
displays no noticeable sexual dimorphism, except in _S. sordida_, in
which there are sexual differences and geographic variation (see p.
324).

_Hands and Feet_

The characters of the hands and feet are among the most taxonomically
important external features in _Smilisca_. Consistent differences exist
in relative lengths of the digits, size of subarticular tubercles, size
and number of supernumerary tubercles, size and shape of the inner
metatarsal tubercle, and amount of webbing (Pls. 4 and 5). In the
_baudini_ group the series of species (_baudini-phaeota-cyanosticta_)
show a progressive increase in amount of webbing in the hand and a
decrease in number, and corresponding increase in size, of supernumerary
tubercles. The amount of webbing in the feet of _S. baudini_ and
_phaeota_ is about the same, but the webbing is slightly more extensive
in _S. cyanosticta_. _Smilisca puma_ is unique in the genus in lacking
webbing in the hand; furthermore, this species is distinctive in having
many large subarticular tubercles on the hand and a relatively small
inner metatarsal tubercle. The two stream-inhabitants, _S. sila_ and
_sordida_, have shorter and stouter fingers than the other species. The
webbing is most extensive in both the hands and feet of these species,
which also are distinctive in having many small supernumerary tubercles
on the feet.

_Ontogenetic Changes_

Minor ontogenetic changes in structure involve the shape of the snout,
relative size of the eye, development of the tympanum, and amount of
webbing in the hand. In recently metamorphosed young the snout is more
rounded than in adults; the canthus and loreal concavity are not
evident. Usually the tympanum is not differentiated in recently
metamorphosed young, and the eye is proportionately large. The webbing
in the feet is completely developed at metamorphosis, but young
individuals have noticeably less webbing in the hand than do adults of
the same species.


Coloration

Some of the most distinctive characters of the species of _Smilisca_ are
color and pattern of the living frogs. Although many chromatic features
are lost or subdued in preserved specimens, the patterns usually
persist.

_Metachrosis_

Change in color, well known in frogs, is common in hylids, especially in
species having green dorsal surfaces (_Phyllomedusa_ is a notable
exception). The non-green _Smilisca_ (_puma_, _sila_, and _sordida_)
changes color, but this mostly is a change in intensity of color. In
these species the markings usually are most distinct at night;
frequently by day the frogs become pallid. The most striking examples of
metachrosis in _Smilisca_ are found in the _baudini_ group, in which the
dorsal ground-color changes from green to tan; correlated with the
change in ground-color may be a corresponding change in the dorsal
markings, but the dorsal markings may change to the opposite color.


Chromosomes

Chromosomes of all six species of _Smilisca_ were studied by means of
the propriono-orcein squash technique described by Duellman and Cole
(1965). Karyotype analysis was attempted for several species by means of
intraperitoneal injections of colchicine, which affected the mitotic
cells as desired, but the testes examined contained too few mitotic
cells to allow accurate determination of karyotypes.

Haploid (_n_) chromosome numbers were determined from cells in
diakinesis, metaphase I, and metaphase II of meiosis. Diploid (2_n_)
chromosome numbers were determined from cells in late prophase and
metaphase of mitosis. Chromosome counts from as few as 23 meiotic cells
of _S. phaeota_ and as many as 80 cells of _S. sordida_ reveal a
constant haploid (_n_) number of 12; counts of chromosomes in one to
five mitotic cells in all species, except _S. sila_, reveal that the
diploid (2_n_) number is 24.



NATURAL HISTORY


Breeding

Like most hylid frogs _Smilisca_ is most readily collected and observed
when individuals congregate for breeding.

_Time of Breeding_

_Smilisca_ breeds primarily in quiet water and reaches its height of
breeding activity at times of plentiful rainfall,--usually from May
through October. Through most of its range _Smilisca baudini_ breeds in
those months, but in some places where abundant rain falls in other
seasons, the species breeds at those times. For example, in southern El
Petén and northern Alta Verapaz, Guatemala, _Smilisca baudini_ has been
found breeding in February and March. The other pond-breeding species
(_S. cyanosticta_, _phaeota_, and _puma_) live in regions lacking a
prolonged dry season, and possibly they breed throughout the year, but
breeding activity seems to be greatest in the rainiest months.

The two stream-breeding species (_S. sila_ and _sordida_) breed in the
dry season when the streams are low and clear, principally in December
through April. At high elevations the species sometimes breed in the
rainy season; also, individuals sometimes breed in the short dry season
(summer canicula) in July and August.

At several localities species have been found breeding at different
times of the year: _S. baudini_ in March and July at Chinajá, Guatemala;
_S. phaeota_ in April and August at Palmar Sur, Costa Rica; _S. puma_ in
February and July at Puerto Viejo, Costa Rica; and _S. sila_ in
February, April, and August at El Volcan, Panamá. These observations
indicate only that the population breeds at more than one time in the
year, but do not provide any evidence on the breeding cycles of the
individual frogs. This is one important aspect of the natural history of
_Smilisca_ for which we lack data.

_Breeding Sites_

All members of the genus _Smilisca_ presumably deposit their eggs in
water.

_Smilisca baudini_ usually breeds in temporary rain pools; often these
are nothing more than shallow, muddy puddles. In other instances the
sites are extensive ditches or large flooded areas (Pl. 8, Fig. 1). This
species is an opportunistic breeder, and males gather at any of a wide
variety of suitable breeding sites that are formed by torrential rains
in the early part of the rainy season. _Smilisca baudini_ nearly always
breeds in open pools having bare earthen edges. Frequently congregations
of _S. baudini_ are found at such small pools, but are absent from
nearby large ponds surrounded by vegetation.

Little is known of the breeding habits of _S. cyanosticta_, which
inhabits humid forests on foothills and lowlands. Apparently its
breeding sites are not unlike those of _S. phaeota_, which usually are
pools surrounded by vegetation (Pl. 8, Fig. 2), although sometimes males
of _S. cyanosticta_ call from open muddy puddles. In uplands, where
standing water is uncommon, this species breeds in quiet pools in
streams.

_Smilisca puma_ breeds in grass-choked ponds and marshes, where the
males call from bases of dense clumps of grass in the water (Pl. 9, Fig.
1).

_Smilisca sila_ and _S. sordida_ differ noticeably from other species in
the genus by breeding exclusively in streams, where males usually call
from rocks or gravel bars in or at the edges of streams (Pl. 9, Fig. 2);
sometimes individuals perch on bushes overhanging streams. In the
streams, or parts of streams, utilized by these frogs the water is
clear, shallow, and has a slow gradient; occasional males have been
found calling along cascading mountain streams.

Breeding choruses composed of ten or more species of frogs are not
uncommon in Middle America, but _Smilisca_ usually breeds alone or with
one or two other species and at the most five others. This tendency
towards solitary breeding possibly is the result of selection of
breeding sites that are unsuitable to many other species of frogs.
Nevertheless, many other species of frogs have been found at the
breeding sites with the various species of _Smilisca_; these breeding
associates (Table 7) are most numerous for _S. baudini_, which has a
broad geographic range, including a variety of habitats.

_Breeding Behavior_

_Calling sites._--All species of _Smilisca_ usually call from the
ground, including rocks and gravel bars; some individuals sit in shallow
water near the edge of the pool or stream. Sometimes males of _S.
baudini_, _sila_, and _sordida_ call from low bushes or trees near the
breeding site. One _S. baudini_ was observed calling while it was
floating on the surface of a pond. _Smilisca cyanosticta_, _phaeota_,
and _puma_ call from secluded places at the edge of the water or in the
water, whereas _S. baudini_, _sila_ and _sordida_ call from open
situations.


  Table 7.--Breeding Associates of the Various Species of Smilisca.

  ==============================+========+============+========+=====+=====+========
  Associate                     |_S.     |_S.         |_S.     |_S.  |_S.  |_S.
                                |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_
  ------------------------------+--------+------------+--------+-----+-----+--------
  _Rhinophrynus dorsalis_       |   X    |     -      |   -    |  -  |  -  |   -
  _Leptodactylus bolivianus_    |   -    |     -      |   X    |  -  |  -  |   -
  _Leptodactylus labialis_      |   X    |     -      |   X    |  -  |  -  |   -
  _Leptodactylus melanonotus_   |   X    |     -      |   X    |  X  |  X  |   -
  _Leptodactylus occidentalis_  |   X    |     -      |   -    |  -  |  -  |   -
  _Leptodactylus quadrivittatus_|   -    |     -      |   X    |  -  |  -  |   -
  _Leptodactylus pentadactylus_ |   -    |     -      |   X    |  X  |  -  |   X
  _Engystomops pustulosus_      |   X    |     -      |   X    |  -  |  -  |   -
  _Bufo canaliferus_            |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo cavifrons_              |   -    |     X      |   -    |  -  |  -  |   -
  _Bufo coccifer_               |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo coniferus_              |   -    |     -      |   X    |  -  |  -  |   -
  _Bufo cristatus_              |   -    |     X      |   -    |  -  |  -  |   -
  _Bufo gemmifer_               |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo haematiticus_           |   -    |     -      |   X    |  -  |  X  |   X
  _Bufo kellogi_                |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo luetkeni_               |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo marinus_                |   X    |     -      |   X    |  X  |  X  |   X
  _Bufo marmoreus_              |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo mazatlanensis_          |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo melanochloris_          |   -    |     -      |   X    |  -  |  X  |   X
  _Bufo perplexus_              |   X    |     -      |   -    |  -  |  -  |   -
  _Bufo typhonius_              |   -    |     -      |   X    |  -  |  X  |   -
  _Atelopus varius_             |   -    |     -      |   -    |  -  |  X  |   X
  _Diaglena reticulata_         |   X    |     -      |   -    |  -  |  -  |   -
  _Diaglena spatulata_          |   X    |     -      |   -    |  -  |  -  |   -
  ------------------------------+--------+------------+--------+-----+-----+--------

  Table 7.--_Continued_
  ==============================+========+============+========+=====+=====+========
  Associate                     |_S.     |_S.         |_S.     |_S.  |_S.  |_S.
                                |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_
  ------------------------------+--------+------------+--------+-----+-----+--------
  _Hyla boulengeri_             |   -    |     -      |   X    |  -  |  -  |   -
  _Hyla colymba_                |   -    |     -      |   -    |  -  |  X  |   -
  _Hyla ebraccata_              |   X    |     -      |   X    |  -  |  -  |   -
  _Hyla elaeochroa_             |   -    |     -      |   X    |  X  |  -  |   -
  _Hyla eximia_                 |   X    |     -      |   -    |  -  |  -  |   -
  _Hyla legleri_                |   -    |     -      |   -    |  -  |  -  |   X
  _Hyla microcephala_           |   X    |     -      |   X    |  -  |  -  |   -
  _Hyla phlebodes_              |   -    |     -      |   X    |  X  |  -  |   -
  _Hyla picta_                  |   X    |     -      |   -    |  -  |  -  |   -
  _Hyla robertmertensi_         |   X    |     -      |   -    |  -  |  -  |   -
  _Hyla rosenbergi_             |   -    |     -      |   X    |  -  |  -  |   -
  _Hyla rufioculis_             |   -    |     -      |   -    |  -  |  -  |   X
  _Hyla smithi_                 |   X    |     -      |   -    |  -  |  -  |   -
  _Hyla staufferi_              |   X    |     -      |   -    |  -  |  -  |   -
  _Hyla walkeri_                |   X    |     -      |   -    |  -  |  -  |   -
  _Phrynohyas inflata_          |   X    |     -      |   -    |  -  |  -  |   -
  _Phrynohyas spilomma_         |   X    |     -      |   -    |  -  |  -  |   -
  _Phrynohyas venulosa_         |   X    |     -      |   -    |  -  |  -  |   -
  _Phyllomedusa callidryas_     |   X    |     -      |   X    |  -  |  -  |   -
  _Phyllomedusa dacnicolor_     |   X    |     -      |   -    |  -  |  -  |   -
  _Phyllomedusa moreleti_       |   X    |     X      |   -    |  -  |  -  |   -
  _Pternohyla fodiens_          |   X    |     -      |   -    |  -  |  -  |   -
  _Smilisca baudini_            |   X    |     X      |   -    |  -  |  -  |   -
  _Smilisca cyanosticta_        |   X    |     X      |   -    |  -  |  -  |   -
  _Smilisca phaeota_            |   -    |     -      |   X    |  -  |  -  |   -
  _Smilisca puma_               |   -    |     -      |   -    |  X  |  -  |   -
  ------------------------------+--------+------------+--------+-----+-----+--------

  Table 7.--_Concluded_
  ==============================+========+============+========+=====+=====+========
  Associate                     |_S.     |_S.         |_S.     |_S.  |_S.  |_S.
                                |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_
  ------------------------------+--------+------------+--------+-----+-----+--------
  _Smilisca sila_               |   -    |     -      |   -    |  -  |  X  |   X
  _Smilisca sordida_            |   -    |     -      |   X    |  -  |  X  |   X
  _Triprion petasatus_          |   X    |     -      |   -    |  -  |  -  |   -
  _Cochranella fleischmanni_    |   -    |     -      |   -    |  -  |  X  |   X
  _Centrolene prosoblepon_      |   -    |     -      |   -    |  -  |  X  |   -
  _Gastrophryne elegans_        |   X    |     -      |   -    |  -  |  -  |   -
  _Gastrophryne olivacea_       |   X    |     -      |   -    |  -  |  -  |   -
  _Gastrophryne usta_           |   X    |     -      |   -    |  -  |  -  |   -
  _Hypopachus alboventer_       |   X    |     -      |   -    |  -  |  -  |   -
  _Hypopachus caprimimus_       |   X    |     -      |   -    |  -  |  -  |   -
  _Hypopachus inguinalis_       |   X    |     -      |   -    |  -  |  -  |   -
  _Hypopachus maculatus_        |   X    |     -      |   -    |  -  |  -  |   -
  _Hypopachus oxyrrhinus_       |   X    |     -      |   -    |  -  |  -  |   -
  _Hypopachus variolosus_       |   X    |     -      |   -    |  -  |  -  |   -
  _Rana palmipes_               |   X    |     -      |   X    |  X  |  -  |   -
  _Rana pipiens_                |   X    |     -      |   -    |  -  |  -  |   -
  _Rana warschewitschi_         |   -    |     -      |   X    |  -  |  X  |   X
  ------------------------------+--------+------------+--------+-----+-----+--------


_Chorus structure._--Limited observations on some of the species of
_Smilisca_ show a definite organization of the calling behavior of
individuals. _Smilisca baudini_ and _S. phaeota_ call in duets. This is
especially noticeable in _S. baudini_, in which the members of a duet
often call from sites separated by only a few centimeters. The call of
_S. baudini_ consists of a series of like notes (see description of call
in following section); the duration of each note is about equal to the
interval between notes. Normally one individual utters one note, pauses,
and utters a single note again, or series of two or three notes. If
there is no response, the first individual often waits several seconds
or even several minutes and then repeats the call. The second individual
usually responds after the first or second note of the sequence. The
notes of the second individual usually are spaced so that they are
emitted in the intervals between the notes of the first individual. This
can be shown diagrammatically by having the figure "1" represent notes
of the first individual and figure "2," the notes of the second; an
empty interval is represented by "0":

   1-0-1-2-1-2-1-2-1-2-1-2

Usually a chorus is initiated by one duet and is quickly picked up by
other individuals also calling in duets. A numerical representation of a
chorus of eight frogs would approximate the following organization:

   1-0-1-2-1-2-1-2-1-2-1-2-1-2
           3-0-3-4-3-4-3-4-3-4-3-4-3
            5-6-5-6-5-6-5-6-5-6-5-6
            7-8-7-8-7-8-7-8-7-8-7-8

After the first one or two duets are initiated, the second individuals
in the following duets usually call immediately after their respective
partners have given the first notes. The other noteworthy aspect about
the organization is that the entire chorus usually stops abruptly.
Normally the first duet stops calling shortly before the others, but
this is not invariable. Often one duet or one individual will emit
several notes after the rest of the frogs have become silent. An
interval of several minutes sometimes elapses before the chorus begins
again. Successive choruses apparently are initiated by the same duet.
Responses can be initiated artificially by imitating the call, and
sometimes any loud noise will start a chorus.

Similar duets have been observed in _S. phaeota_. In this species the
intervals are often much longer than the notes, and if two males are
calling in close proximity, their calls can be mistaken for those of one
individual. _Smilisca phaeota_ does not congregate in large numbers;
usually only two males call from one restricted site.

_Smilisca sila_ has a call consisting of a primary note followed by one
or more secondary notes. Males often call in duets, but not necessarily
so. In a duet, the first male usually utters only primary notes until
the second individual responds; then each individual produces a rapid
series of secondary notes.

_Smilisca puma_ also produces primary and secondary notes. Although
individuals sometimes call alone, duets, trios, or quartets were more
common. The chorus is initiated by one individual uttering primary notes
until joined by the second, third, and fourth frogs. In one quartet in a
marsh 7.5 kilometers west of Puerto Viejo, Costa Rica, on February 19,
1965, the same individual initiated four consecutive choruses. Each time
the second member of the chorus was the same; the third and fourth frogs
joined the chorus nearly simultaneously.

Individuals of _S. sordida_ are usually irregularly situated along a
stream. No duets or other combinations of individuals are apparent in
the chorus structure, but once an individual calls, a frog nearby calls
almost immediately; then a frog near the second individual calls, and so
on. The resulting series of calls gives the impression that the sound is
moving along the stream as successive individuals join the chorus and
the first callers become quiet. It is not known if the same individual
initiates successive choruses or if the order of calling is the same in
subsequent choruses.

These limited observations on chorus structure in _Smilisca_ show the
presence of behavioral organization. The methods of establishing the
organization and the significance of the call-order in breeding have yet
to be discovered.

Calling males of _S. baudini_ are often close together; some individuals
have been observed almost touching one another, but no indication of
territoriality or aggressive behavior has been witnessed. The more
distant spacing of the stream-breeding species _S. sila_ and _S.
sordida_ may be a function of calling-territories, but no direct
evidence is available to substantiate this supposition.

_Sex recognition and amplexus._--Observations on _Smilisca baudini_
indicate that the calls of males attract females. At Tehuantepec,
Oaxaca, México, a female was first observed about two meters away from a
male calling at the edge of a rain pool; in a series of short hops she
progressed directly towards the male, although vegetation obscured him
until she was less than a meter away. When she approached to within
about 20 centimeters of the male, he took notice of her, moved to her,
and clasped her. At Chinajá, Alta Verapaz, Guatemala, a female swam
directly across a pool about three meters wide to a calling male. Her
line of movement took her within a few centimeters of a silent male, to
whom she paid no attention. She stopped just in front of the calling
male, which immediately clasped her. At a large muddy pond 4 kilometers
west-northwest of Esparta, Puntarenas, Costa Rica, a female was observed
swimming toward a small submerged tree; a male was calling from a branch
about one meter above the water. The female climbed to a branch about 20
centimeters below the male, which upon seeing her there immediately
jumped down and clasped her. These few observations of _S. baudini_ show
that in this species females are capable of locating calling males by
means of phono-orientation; visual reception on the part of females
seems to be secondary. Contrariwise, males apparently become aware of
the proximity of females by seeing them; once a male sees a female he
usually tries to clasp her. Possibly the males receive stimuli by means
of chemo-reception, but in each observed instance the male obviously
looked at the female.

Amplexus is axillary in all members of the genus. Normally amplexing
males hunch their backs and press their chins to the females' backs.
Clasping pairs are usually found at the edge of the water, but sometimes
amplexus takes place in trees or bushes.

_Egg deposition._--Oviposition has been observed only in _Smilisca
baudini_. On the night of June 28, 1961, at Chinajá, Alta Verapaz,
Guatemala, a clasping pair was observed at the edge of a shallow rain
pool. After sitting for several minutes in shallow water, the female
(with male on her back) swam part way across the pool and grasped an
emergent stick with one hand. The female's body was nearly level with
the surface of the water, and her hind legs were outstretched as
deposition commenced; eggs were extruded rapidly. After a few seconds
the female moved slowly to another twig a few centimeters away and
deposited more eggs. This process was repeated until the female was
spent. The spawn resulted in a surface film covering roughly one square
meter. It is doubtful if this type of egg deposition occurs in any other
species in the genus, especially those that lay their eggs in streams.

_Breeding Call_

The breeding calls of the six species of _Smilisca_ are alike in their
explosive nature. Calls are emitted quickly with a short burst of air
filling the vocal sac, which immediately deflates. Phonetically the
calls can be described as a single "wonk" or series of such notes in _S.
baudini_ and _S. cyanosticta_, a low growl in _S. phaeota_, a relatively
high pitched rattle in _S. sordida_, and a low squawk usually followed
by one or more rattling secondary notes in _S. puma_ and _S. sila_.
Quantitatively, the calls of the six species differ in number of notes,
duration of notes, and in pitch (Table 8, Pls. 10 and 11). Although no
measurements were taken on the intensity of the calls, we observed in
the field that each of the species has a loud voice. The call of _S.
baudini_ seems to carry farther than any of the others.


  Table 8.--Comparison of Breeding Calls in Smilisca. (Observed Range
     Given in Parentheses Below Mean. In Species Having Primary and
     Secondary Notes, Only the Primary Notes Are Analyzed Here.)

  ==============+====+=======+===========+=========+=========+=======================+
                |    | Notes |           |         | Funda-  |         Major         |
     Species    |    |  per  |  Duration | Pulses  | mental  |   frequencies (cps)   |
                | N  |  call |  of note  |  per    |frequency+-----------+-----------+
                |    | group | (seconds) | second  | (cps)   |   Lower   |   Upper   |
  --------------+----+-------+-----------+---------+---------+-----------+-----------+
  _S.           | 20 |  8.0  |    0.11   |  174.7  |  166.2  |    351    |   2507    |
    baudini_    |    | (2-15)|(0.09-0.13)|(140-195)|(135-190)| (175-495) |(2400-2725)|
                |    |       |           |         |         |           |           |
  _S.           | 10 |  1.2  |    0.38   |  147.0  |  145.1  |    841    |   1894    |
    cyanosticta_|    | (1-2) |(0.25-0.45)|(110-180)|(135-160)| (480-975) |(1600-2100)|
                |    |       |           |         |         |           |           |
  _S. phaeota_  | 10 |  1.6  |    0.31   |  116.0  |  143.0  |    372    |    --     |
                |    | (1-2) |(0.10-0.45)|(100-130)|(110-165)| (330-495) |    --     |
                |    |       |           |         |         |           |           |
  _S. puma_     | 28 |  3.7  |    0.13   |  208.2  |  145.6  |    743    |   1868    |
                |    | (2-10)|(0.06-0.35)|(187-240)|(125-200)| (495-980) |(1456-2240)|
                |    |       |           |         |         |           |           |
  _S. sila_     | 15 |  2.4  |    0.16   |  108.5  |  103.0  |    899    |   2218    |
                |    | (1-6) |(0.06-0.28)| (97-120)| (90-115)| (665-1180)|(1980-2700)|
                |    |       |           |         |         |           |           |
  _S. sordida_  | 19 |  1.7  |    0.29   |  104.7  |  123.1  |   1216    |   2694    |
                |    | (1-6) |(0.18-0.45)| (78-135)| (90-140)|(1150-1540)|(2340-2990)|
  --------------+----+-------+-----------+---------+---------+-----------+-----------+


_Call rate._--The rate at which call-groups are produced varies from one
every few seconds to one in several minutes. In _S. baudini_,
_cyanosticta_, _phaeota_, and _sordida_, call-groups are produced as
frequently as every 12 seconds, but usually more time elapses between
call groups. In _S. sordida_, five or more minutes sometimes elapse
between call-groups. The interval is somewhat less in _S. phaeota_.
Calls are repeated at much shorter intervals in _S. puma_ (5-55 seconds)
and _S. sila_ (4-20 seconds).

_Notes per call-group._--Except for _S. puma_ and _S. sila_, the series
of notes produced in any given call of a species of _Smilisca_ is
essentially the same; there is no differentiation into primary and
secondary notes. _Smilisca cyanosticta_ and _S. phaeota_ emit only one
or two relatively long notes per call-group, whereas _S. baudini_ and
_S. sordida_ produce as many as 15 and 6 notes, respectively. Males of
_S. puma_ and _S. sila_ often produce only the primary note; sometimes
this is done several times before the secondary notes are produced. For
example, one _S. puma_ (KU 91711; tape No. 379) produced the following
number of notes in consecutive call-groups: 1, 1, 1, 1, 2, 2, 3, 1, 4;
secondary notes are present in only four of the nine call-groups. A
typical series of consecutive call-groups in _S. sila_ (KU 91852; Tape
No. 385) has 1, 1, 1, 2, 4, 2 notes per call-group; secondary notes are
present in only half of the call-groups. _Smilisca puma_ apparently
always produces at least two primary notes before emitting secondary
notes; sometimes only primary notes are produced in one series of calls.
The number of secondary notes following a given primary varies from one
to nine; the modal number is one, and the mean is three in 27
call-groups. _Smilisca sila_ frequently begins a series of calls with
two or more primary notes, but sometimes the first primary note is
followed immediately by two or more secondary notes. The number of
secondary notes following a given primary varies from one to five; the
modal number is one, and the average is two in 13 call-groups.

_Duration._--The average duration of call-groups consisting of two or
more notes is 1.18 seconds in _S. baudini_; 1.02 in _cyanosticta_, 0.91
in _phaeota_, 1.32 in _puma_, 1.48 in _sila_, and 1.29 in _sordida_.
Although there is considerable variation in the lengths of the notes
(only primary notes in _S. puma_ and _sila_ are considered here), _S.
cyanosticta_, _phaeota_, and _sordida_ have noticeably longer notes than
do the other species (Table 8). The secondary notes are longer than the
primary notes in _S. puma_ (average 0.27 secs. as compared with 0.13
secs.) and in _S. sila_ (average 0.25 secs., as compared with 0.16
secs.).

_Note repetition rate._--The rate at which notes in call-groups
containing two or more notes are produced varies in _S. baudini_ from
2.5 to 7.1 (average, 3.7) calls per second; _cyanosticta_, 1.8-2.1
(1.9); _phaeota_, 2.0-2.4 (2.2); _puma_, 1.9-2.9 (2.2); _sila_, 1.3-2.4
(1.8); and _sordida_, 1.5-2.6 (2.1). _Smilisca baudini_, which has
notes of short duration (0.09 to 0.13 seconds), has the fastest
note-repetition rate. Although the individual notes of _S. cyanosticta_
and _S. phaeota_ are relatively long (average, 0.38 and 0.31 seconds,
respectively), the intervals between the notes is short; consequently,
their note-repetition rates do not differ greatly from those of _S.
puma_ and _S. sila_, which have shorter notes (average, 0.13 and 0.16
seconds, respectively) but longer intervals between notes.

_Pulse rate._--Pulses vary in frequency from 78 to 240 per second in the
calls analyzed (only primary notes in _S. puma_ and _S. sila_), but the
variation in any given species is much less than that in the entire
genus (Table 8). _Smilisca puma_ is outstanding in having a high pulse
rate, which is approached only by that of _S. baudini_. Even in the
species having the lowest pulse rates, the pulsations are not audible.
The secondary notes produced by _S. puma_ and _S. sila_ have a slower
pulse rate than the primary notes; often the pulses are audible. In _S.
puma_ the pulse rate of secondary notes is sometimes as low as 48 pulses
per second, and in _S. sila_ still lower (as low as 40 pulses per
second). The upper limits of pulse rate in the secondary notes in these
species merge imperceptibly with the rates of the primary note;
consequently, on the basis of pulse rate alone it is not always possible
to distinguish primary from secondary notes.

_Frequency._--_Smilisca_ produces noisy (as opposed to more musical)
calls, and the energy is distributed throughout the frequency spectrum;
the calls are poorly modulated, except in _S. sordida_, in which two
usually discrete bands of frequency are present (Pl. 11C). For the most
part the calls of _Smilisca_ consist of little modified energy of the
fundamental frequency and of its harmonics, some of which are
emphasized.

The upper frequency range varies within each species and even within the
calls of one individual. _Smilisca phaeota_ has the lowest upper
frequencies; no calls ranged above 4400 cycles per second (cps.), and
half of the calls never exceeded 3000 cps. _Smilisca cyanosticta_
produces calls in which the upper frequency is below 7000 cps. and
usually below 6000 cps. Likewise, _S. puma_ produces calls that are
below 7000 cps., whereas _S. sila_ has frequencies of up to 8400 cps. In
both _S. baudini_ and _S. sordida_, the highest frequencies attained are
about 9100 cps. Variation in the highest frequencies in a series of
consecutive calls by one individual frog was noted in all species. Such
variation is especially prevalent in _S. puma_; for example one
individual (KU 87771; Tape No. 376) recorded at a temperature of 24° C.
at 7.5 kilometers west of Puerto Viejo, Heredia Province, Costa Rica, on
July 31, 1964, produced three consecutive primary notes having upper
frequencies of about 6000, 4000, and 4000 cps., respectively. Apparently
in a given species the production of the higher frequencies in some
notes and not in others is correlated with the amount of distention of
the vocal sac and is not dependent upon the structure or tension of the
vocal cords.

Although the dominant frequency in _S. sordida_ is lower than that in
_S. baudini_ and _S. cyanosticta_, the call of the former is audibly
higher-pitched. This is due primarily to the emphasis on certain
harmonics at a high frequency (sometimes as high as 9000 cps.) in _S.
sordida_, whereas in _S. baudini_ and other species, if harmonics are
present at those frequencies, they are not emphasized.

The fundamental frequencies are as low as 90 cps. in _S. sila_ and _S.
sordida_ and as high as 200 cps. in _S. puma_ (Table 8). The fundamental
frequency seemingly is relatively unimportant in determining the general
pitch of the call, a characteristic most dependent on the dominant
frequency and emphasized harmonics in the higher-frequency spectrum. In
none of the species is the fundamental the dominant frequency. In the
low-pitched call of _S. phaeota_ the dominant frequency is the third
harmonic (the second harmonic above the fundamental frequency, which is
the first harmonic). In all other species a much higher harmonic is
dominant; for examples, in _S. cyanosticta_ harmonics from 10 to 15 are
dominant; in _S. baudini_, 15-19; and _S. sila_, 20-30.

A glance at the audiospectrographs and their accompanying sections (Pls.
10 and 11) reveals the presence of two emphasized bands of frequency in
all species except _S. phaeota_, in which only the lower band is
present. These two bands of emphasized harmonics are part of a
continuous, or nearly continuous, spread of energy throughout the
frequency spectrum, except in _S. sordida_ in which the bands are
usually distinct. As shown in the sections, certain harmonics in each of
the bands are emphasized with nearly equal intensity. Therefore, with
the exception of _S. phaeota_, the calls of _Smilisca_ are characterized
by two major frequencies, one of which is the dominant frequency and the
other is a subdominant frequency (Table 8). The upper major frequency is
dominant in all calls in _S. baudini_ and _S. cyanosticta_, but either
major frequency may be dominant in other species. The upper major
frequency is dominant in 65 per cent of calls by _S. puma_, 87 per cent
in _S. sila_, and 68 per cent in _S. sordida_. Individuals of these
three species sometimes produce a series of calls in which the dominant
frequency changes from one of the major frequencies to the other. Four
consecutive notes emitted by an individual of _S. sordida_ recorded 13
kilometers east-northeast of Golfito, Puntarenas Province, Costa Rica,
had dominant frequencies of 910, 1950, and 750 cps., respectively. In
each case, an alternation of major frequencies took place in respect to
dominance. An individual of _S. puma_ from 7.5 kilometers west of Puerto
Viejo, Costa Rica, produced a primary note followed by one secondary
note; each note had major frequencies at 600 and 1800 cps.; the dominant
frequency of the primary note was at 1800 cps., whereas in the secondary
note the dominant frequency was at 600 cps. The difference in emphasis
on the major frequencies is so slight that shift in dominance is not
audible.

_Effect of temperature on calls._--The present data are insufficient to
test statistically the correlation between temperature and variation
within certain components of the calls in _Smilisca_, but even a crude
graph shows some general correlations. The widest range of temperatures
is associated with the recordings of _S. baudini_. Three individuals
recorded at a temperature of 30° C. at Tehuantepec, Oaxaca, had pulse
rates of 180 pulses per second and fundamental frequencies of 160-180
cps., as compared with an individual recorded at a temperature of 17°
C., which had a pulse rate of 140 and a fundamental frequency of 135
cps. All individuals of _S. baudini_ recorded at higher temperatures had
faster pulse rates and higher fundamental frequencies. Pulse rates
differ in the other species in the genus but less strikingly (probably
owing to narrower ranges of temperatures at which recordings were made).
In five recordings of _S. sordida_ made at 20° C. the pulse rate is
80-90, as compared with four recordings made at 25° C. having pulse
rates of 120-135. Thirteen recordings of _S. sila_ made at 17° C. have
pulse rates of 97-112 (average 105); one individual recorded at 26° C.
has 120 pulses per second. Seemingly no correlation exists between
temperature and other characteristics of the calls, such as duration and
rate of note-repetition.

_The breeding call as an isolating mechanism._--Blair (1958), Bogert
(1960), Duellman (1963a), Fouquette (1960), Johnson (1959), and others
have provided evidence that the breeding calls of male hylids (and
other anurans) serve as isolating mechanisms in sympatric species. In
summarizing this discussion of the breeding calls of _Smilisca_ we want
to point out what seem to be important differences in the calls that may
prevent interspecific hybridization in sympatric species of _Smilisca_.

The genus is readily divided into two species-groups on morphological
characters; this division is supported by the breeding calls. In the
species of the _baudini_ group the calls are unmodulated and lack
secondary notes. In the _sordida_ group the calls either have secondary
notes or are modulated.

_Smilisca baudini_ occurs sympatrically with _S. cyanosticta_ and _S.
phaeota_; where they occur together, both species sometimes breed in
like places at the same time. We are not aware of these species breeding
synchronously at exactly the same site, although _S. baudini_ and _S.
cyanosticta_ were calling on the same nights and less than 100 meters
apart in Oaxaca in June, 1964. Regardless of their respective breeding
habits, sympatric species have calls that differ notably. Except for the
higher fundamental and dominant frequencies, the calls of _S.
cyanosticta_ and _S. phaeota_ closely resemble one another, but the
calls of both species differ markedly from that of _S. baudini_. The
geographic ranges of _S. cyanosticta_ and _S. phaeota_ are widely
separated.

The calls of the allopatric species _S. puma_ and _S. sila_ are not
greatly different. _Smilisca sordida_ has a distinctive call and occurs
sympatrically with _S. puma_ and _S. sila_. In the streams in southern
Costa Rica _S. sordida_ and _S. sila_ breed synchronously, but the
high-pitched modulated call of the former is notably different from the
lower, unmodulated call of _S. sila_.

The data indicate that the calls of related sympatric species differ
more than the calls of related allopatric species. We postulate that
these differences evolved to support the reproductive isolation of the
sympatric species. The data are insufficient to determine geographic
variation in the calls and to determine if differences in the calls are
enhanced in areas of sympatry as compared with the allopatric parts of
the ranges.

_Other calls._--As stated previously, there is no direct evidence of
territoriality in _Smilisca_; we have heard no calls that can be
definitely identified as territorial. Single notes of _S. baudini_,
_phaeota_, and _sila_ have been heard by day, just prior to rains, or
during, or immediately after rains. Such calls can be interpreted as
"rain calls," which are well known in _Hyla eximia_ and _Hyla
squirella_. Distress calls are known in several species of _Rana_ and in
_Leptodactylus pentadactylus_; such calls result from the rapid
expulsion of air over the vocal cords and with the mouth open. Distress
calls have been heard from _S. baudini_. At Charapendo, Michoacán,
México, a male that had one hind limb engulfed by a _Leptodeira
maculata_ emitted several long, high-pitched cries. A clasping pair of
_S. baudini_ was found in a bush at the edge of a marshy stream 2
kilometers northeast of Las Cañas, Guanacaste Province, Costa Rica. When
the pair was grasped, the female emitted a distress call.


Eggs

Eggs of _S. baudini_, _cyanosticta_, and _phaeota_ have been found in
the field, and eggs of _S. sila_ have been observed in the laboratory.
The eggs of _S. puma_ and _sordida_ are unknown. Insofar as known,
_Smilisca baudini_ is unique in the genus in depositing the eggs in a
surface film. Each egg is encased in a vitelline membrane, but
individual outer envelopes are lacking. The eggs are small; the diameter
of recently-deposited eggs is about 1.3 mm. and that of the vitelline
membrane is about 1.5 mm. The eggs of _S. cyanosticta_ and _phaeota_ are
deposited in clumps, and the eggs are larger than those of _S. baudini_.
Diameters of eggs of _S. cyanosticta_ are about 2.3 mm., and those of
the outer envelopes are about 4.0 mm. Artificially fertilized eggs of
_S. sila_ raised in the laboratory have diameters of about 2.4 mm.; the
diameter of the outer envelopes is about 4.9 mm.

In order to determine the reproductive potential of the six species,
ovulated eggs were removed from females and counted. The numbers of eggs
recorded are: 3 _S. baudini_--2620, 2940, 3320; 1 _S. cyanosticta_--910;
3 _S. phaeota_--1665, 1870, 2010; 1 _S. puma_--518; 3 _S. sila_--369,
390, 473; 3 _S. sordida_--524, 702, 856. These limited data indicate
that the large species (_S. baudini_, _cyanosticta_, and _phaeota_) have
more eggs than do the smaller species. The stream-breeding species (_S.
sila_ and _sordida_) have relatively few eggs by comparison with the
pond-breeders. Possibly this is a function of size of eggs rather than a
correlation with the site of egg-deposition.


Tadpoles

The acquisition of tadpoles of all of the species of _Smilisca_ has made
possible the use of larval characters in erecting a classification and
in estimating the phylogenetic relations of the several species.
Furthermore, developmental series of tadpoles of four species allow a
comparison of the growth and development in these species. Throughout
the discussion of tadpoles we have referred to the various developmental
stages by the Stage Numbers proposed by Gosner (1960).

_General Structure_

Tadpoles of the genus _Smilisca_ are of a generalized hylid type, having
2/3 tooth-rows, unspecialized beaks, mouth partly or completely bordered
by papillae, lateral fold present in the lips, spiracle sinistral, anal
tube dextral, and caudal musculature extending nearly to tip of caudal
fin. Although minor differences exist in coloration, proportions, and
mouthparts, no great modifications of the basic structure are present.

_Comparison of Species_

The larval characters of the species of _Smilisca_ are compared below
and illustrated in Figures 11-15.

_Shape and Proportions._--The bodies of _S. baudini_, _cyanosticta_,
_phaeota_, and _puma_ are rounded and about as wide as deep; the eyes
are moderately large and directed dorsolaterally, and the nostrils are
about midway between the bluntly rounded snout and the eyes. The mouths
are medium-sized and directed anteroventrally. The bodies of tadpoles of
_S. sila_ and _sordida_ are slightly compressed dorso-ventrally. The
snout is moderately long and sloping; the eyes are larger and directed
more dorsally than in the other species, and the nostrils are closer to
the eyes than the snout. The mouths are moderately large and directed
ventrally.

The tail is about half again as long as the body in _S. baudini_,
_cyanosticta_, _phaeota_, and _puma_; in these species the caudal
musculature is moderately heavy, and the caudal fins are deep. The
caudal musculature is upturned distally in _S. baudini_ and _phaeota_,
and the dorsal fin extends anteriorly onto the body in these two species
and in _S. puma_. The tail is about twice as long as the body in _S.
sila_ and _sordida_. In both species the caudal fins are shallow in
comparison with the depth of the caudal musculature, especially in _S.
sordida_ (Fig. 14); in neither species does the dorsal fin extend
anteriorly onto the body.

    [Illustration: FIG. 11. Tadpoles of _Smilisca baudini_: (A) Stage
       21 (KU 62155) × 10; (B) Stage 25 (KU 68467) × 5; (C) Stage 30
       (KU 60018) × 4; (D) Stage 41 (KU 60018) × 3.]

_Mouthparts._--The mouth of _S. sordida_ is completely bordered by two
rows of papillae, whereas in the other species the median part of the
upper lip is devoid of papillae. _Smilisca baudini_ and _puma_ have two
rows of papillae; _S. sila_ has one complete row (except medially on the
upper lip) and one incomplete row, and _S. cyanosticta_ and _phaeota_
have only one row (Fig. 15). All species have numerous papillae in the
lateral fold; the fewest lateral papillae are found in _S. cyanosticta_
and _phaeota_. Although all species have two rows of teeth in the upper
jaw and three rows in the lower jaw, specific differences in the nature
of the rows exist between certain species. The second upper tooth-row is
narrowly interrupted medially in _S. sila_ and _sordida_ and broadly
interrupted in the other species. The first upper row is strongly arched
in _S. puma_, moderately arched in _S. baudini_ and _sila_, and weakly
arched in the other species. In all species the third lower tooth-row is
the shortest, only slightly so in _S. sila_ and _sordida_, but only
about half the length of the second lower row in _S. puma_.

    [Illustration: FIG. 12. Tadpoles of _Smilisca cyanosticta_:
       (A) Stage 21 (KU 87648) (B) Stage 25 (KU 87651) × 5;
       (C) Stage 30 (KU 87652) × 4; (D) Stage 40 (KU 87650) × 3.]

The beaks are well developed and finely serrate in all species. The
lower, broadly V-shaped, beak is slender in _S. puma_, rather robust in
_S. baudini_ and _sila_, and moderately heavy in the other species. The
lateral processes of the upper beak are shortest in _S. puma_ and
longest in _S. baudini_ and _sordida_. In the latter the inner margin of
the upper beak and lateral process have the form of a shallow S, whereas
in the other species the inner margin of the upper beak forms a
continuous arch with the lateral processes (Fig. 15).

    [Illustration: FIG. 13. Tadpoles of _Smilisca phaeota_: (A) Stage 21
       (KU 68479) × 14; (B) Stage 25 (KU 68480) × 5; (C) Stage 30 (KU
       68482) × 4; (D) Stage 40 (KU 68483) × 3.]

_Coloration._--The tadpoles of _Smilisca_ lack the bright colors or bold
markings characteristic of some hylid tadpoles; even so, the subdued
colors and arrangement of pigments provide some distinctive markings by
which the species can be distinguished from one another. The species
comprising the _baudini_ group (_S. baudini_, _cyanosticta_, and
_phaeota_) are alike in having the body brown or grayish brown dorsally
and transparent with scattered brown pigment ventrally. A cream-colored,
crescent-shaped mark is present on the posterior edge of the body; this
mark is usually most noticeable in _S. baudini_ and least so in _S.
cyanosticta_. Other differences in coloration in members of the
_baudini_ group are relative and subtle. _Smilisca phaeota_ usually is
more pallid than _baudini_, and _cyanosticta_ usually is darker than
_baudini_; both species have larger dark markings on the tail than does
_S. phaeota_. _Smilisca baudini_ has a dark streak on the middle of the
anterior one-fourth of the tail (Figs. 11-13).

_Smilisca puma_ is distinctive in having a grayish brown body and dark
gray reticulations on the tail. _Smilisca sila_ and _sordida_ are
distinctive in having pairs (sometimes interconnected) of dark marks on
the dorsal surfaces of the caudal musculature, and in dorsal view the
tail appears to be marked with dark and pale creamy tan transverse bars.
These dark marks, as well as the small flecks on the tail, are brown in
_S. sila_ and red in _sordida_. _Smilisca sila_ has dark brown flecks on
the dorsal surface of the body and small white flecks laterally; these
markings are absent in _S. sordida_ (Fig. 14).

Descriptions of the coloration of living tadpoles are given in the
accounts of the species.

    [Illustration: FIG. 14. Tadpoles of _Smilisca_; (A) _S. puma_,
       Stage 30 (KU 91807); (B) _S. sila_, Stage 25 (KU 80260);
       _S. sordida_, Stage 30 (KU 68475). All × 3.5.]

_Growth and Development_

Information on the growth and development of Middle American hylids is
scanty. Adequate descriptions have been published for _Phyllomedusa
annae_ (Duellman, 1963b), _Phrynohyas venulosa_ (Zweifel, 1964), and
_Triprion petasatus_ (Duellman and Klaas, 1964). Material is available
for adequate descriptions of the developmental stages of four species of
_Smilisca_ (Tables 9-12, Figs. 11-13). Because none of the tadpoles was
raised from hatching to metamorphosis, the rate of growth and duration
of the larval stages are unknown.

    [Illustration: FIG. 15. Mouthparts of tadpoles of _Smilisca_;
       (A) _S. baudini_ (KU 60018); (B) _S. puma_ (KU 91807);
       (C) _S. cyanosticta_ (KU 87625); (D) _S. sila_ (KU 80620);
       (E) _S. phaeota_ (KU 68482); (F) _S. sordida_ (KU 68475).
       All ×17.]

  Table 9.--Growth and Development of Tadpoles of Smilisca baudini.
     (Means Are Given in Parentheses After the Observed Ranges.)

  ====================================================================
  Stage | N |    Total length   |   Body length    | Tail length
  --------------------------------------------------------------------
    21  |10 |  5.1-5.4   (5.22) | 2.6-2.7   (2.54) | 2.5-2.7   (2.58)
    24  |10 |  6.0-6.5   (6.20) | 2.3-2.6   (2.45) | 3.5-3.9   (3.69)
    25  |10 |  7.2-8.3   (7.78) | 3.0-3.3   (3.14) | 4.2-5.0   (4.64)
    27  |10 | 18.5-21.5 (20.22) | 8.0-9.0   (8.38) |10.4-13.0 (11.84)
    29  |10 | 21.5-24.5 (22.60) | 8.5-10.0  (9.25) |12.5-14.5 (13.35)
    37  | 3 | 28.5-31.0 (30.00) |11.0-12.5 (11.67) |17.5-19.0 (18.00)
    38  |10 | 35.0-37.5 (35.50) |12.0-13.5 (12.80) |21.5-24.0 (22.70)
    40  | 2 | 34.0-37.0 (35.50) |12.5-13.5 (13.00) |21.5-23.5 (22.50)
    41  |10 | 34.0-37.0 (35.50) |12.5-13.5 (13.00) |21.5-23.5 (22.50)
    42  | 3 | 24.0-30.0 (27.00) |12.5-13.0 (12.67) |11.5-17.0 (14.33)
    45  | 6 | 14.0-24.0 (17.58) |12.5-14.0 (13.37) | 1.5-10.0  (4.17)
    46  |23 |      ----         |12.0-15.5 (13.34) |    ----
  --------------------------------------------------------------------


  Table 10.--Growth and Development of Tadpoles of Smilisca
     cyanosticta. (Means Are Given in Parentheses After the
     Observed Ranges.)

  ======================================================================
    Stage | N |   Total length    |   Body length    |   Tail length
  ----------------------------------------------------------------------
    21    | 10|  5.8-6.5   (6.28) | 2.8-3.1   (3.00) | 3.0-3.5   (3.28)
    25    | 10|  7.9-9.2   (8.44) | 2.7-3.2   (2.96) | 4.8-6.0   (5.48)
    30    |  7| 22.5-25.0 (23.50) | 8.5-9.5   (9.00) |14.0-15.5 (14.57)
    36    | 10| 27.0-30.0 (28.75) | 9.5-11.5 (10.80) |17.0-18.5 (17.95)
    42    |  2| 26.0-27.0 (26.50) |      10.00       |16.0-17.0 (16.50)
    46    |  2|        --         |      14.00       |       --
  ----------------------------------------------------------------------


Hatchlings of three species (_S. baudini_, _cyanosticta_, and _phaeota_)
are available. These larvae have non-functional eyes and large oral
suckers. By the time the larvae have developed to stage 21, external
gills are present, the caudal musculature and caudal fin have been
differentiated, and the head is distinguishable from the body. In stage
21 oral suckers and a large amount of yolk are still present.

The developmental data on the four species show no significant
variations; consequently, we will describe the development of only one
species, _Smilisca phaeota_ (Table 11, Figs. 13 and 16).

_Stage 21._--Bulging cream-colored yolk mass, transparent cornea, and
moderately long, unbranched filamentous gills, and oral suckers present;
mouth having irregular papillae on lower lip; teeth and beaks absent;
caudal myomeres distinct; pigmentation uniform over body and caudal
musculature; caudal fin transparent with scattered small flecks.

    [Illustration: FIG. 16. Relative rate of growth in tadpoles of
       _Smilisca phaeota_ as correlated with developmental stages.
       Formulas for the limb bud refer to its length (L) in relation to
       basal diameter (D).]

_Stage 25._--Operculum complete; gills absent; sinistral spiracle
apparently functional; cloacal tail-piece, nasal capsules, and external
nares present; gut partly formed; mouth bordered by single row of
papillae, except medially; small papillae present in lateral fold of
lips; two upper and three lower tooth-rows present, but not fully
developed; beaks apparently fully developed; depth of dorsal and ventral
fins less than depth of caudal musculature: tip of tail upturned;
pigment on body most dense on dorsum and sides; faint, nearly
pigmentless crescent-shaped mark on posterior edge of body;
concentrations of pigment forming small spots on tail.

_Stage 28._--Mouthparts complete; limb bud about half as long as thick;
other structural features and coloration closely resemble those in stage
25.

_Stage 30._--Limb bud approximately twice as long as thick; body as deep
as wide; dorsal fin deepest just posterior to body; ventral fin deeper
than caudal musculature; tail sharply upturned distally; anal tube
dextral; brown pigment sparse on flanks.


  Table 11.--Growth and Development of Tadpoles of Smilisca phaeota.
     (Means Are Given in Parentheses After the Observed Ranges.)

  ====================================================================
  Stage | N |  Total length     |  Body length     | Tail length
  --------------------------------------------------------------------
    15  | 10|      --           | 1.9-2.1   (1.97) |       --
    16  |  8|      --           | 2.0-2.2   (2.07) |       --
    18  |  4|      --           | 2.2-2.6   (2.31) |       --
    21  |  3|  7.9-8.6   (8.21) | 4.1-4.5   (4.31) |  3.8-4.1   (3.92)
    25  | 10|  8.7-10.6  (9.69) | 4.5-4.8   (4.64) |  4.3-6.0   (5.05)
    26  | 11| 12.3-16.1 (14.01) | 4.2-6.3   (5.60) |  6.7-9.8   (8.41)
    27  | 10| 13.0-15.7 (14.28) | 4.9-6.2   (5.40) |  7.7-10.5  (8.88)
    28  | 13| 13.9-20.9 (15.62) | 5.2-8.3   (5.75) |  8.5-12.6  (9.85)
    29  |  8| 17.8-22.3 (19.79) | 6.3-8.4   (7.19) | 11.5-14.0 (12.60)
    30  |  9| 20.3-24.8 (22.85) | 8.1-10.5  (9.32) | 10.5-15.5 (13.53)
    31  |  5| 24.1-28.5 (26.61) | 9.4-11.2 (10.59) | 14.7-17.3 (16.02)
    34  |  5| 24.8-29.4 (27.31) | 9.2-11.6 (10.73) | 15.6-18.5 (16.80)
    36  |  3| 30.0-30.1 (30.07) |10.1-12.2 (11.15) | 18.9-20.0 (19.44)
    37  |  4| 28.9-34.1 (31.75) |11.5-12.4 (11.88) | 17.4-22.5 (19.88)
    38  |  1| 28.98             |12.88             | 16.10
    39  |  2| 35.6-36.9 (36.25) |14.00             | 21.6-22.9 (22.25)
    40  |  2| 32.3-39.8 (36.05) |14.00             | 18.3-21.8 (20.05)
    43  |  2| 21.5-23.0 (22.25) |14.2-14.8 (14.45) |  6.8-8.8   (7.80)
    44  |  4|      --           |14.5-15.6 (15.08) |         --
    46  | 11|      --           |12.7-16.7 (14.26) |         --
  --------------------------------------------------------------------


  Table 12.--Growth and Development of Tadpoles of Smilisca sordida.
     (Means Are Given in Parentheses After the Observed Ranges.)

  =======+===+==================+==================+=================
   Stage | N |   Total length   |    Body length   |    Tail length
  -------+---+------------------+------------------+-----------------
  25     | 6 | 25.5-28.0 (26.1) |  9.0-9.5   (9.3) | 16.2-18.5 (16.7)
  33     | 2 | 28.5-30.0 (29.3) | 10.2-10.5 (10.4) | 18.0-19.8 (18.9)
  36     | 8 | 29.5-34.5 (32.3) | 10.2-11.7 (10.8) | 19.3-23.0 (21.5)
  37     | 7 | 31.6-37.5 (34.6) | 11.0-12.5 (11.5) | 21.6-25.0 (23.2)
  41     | 3 | 33.0-37.2 (35.2) | 11.6-12.2 (11.9) | 21.4-25.2 (23.2)
  43     | 1 |      ----        |      12.4        |      ----
  46     | 9 |      ----        | 13.1-15.7 (14.9) |      ----
  -------+---+------------------+------------------+-----------------


_Stages 34_, _36_, _37_, and _38_.--Stage 34, foot paddle-shaped with
four toe buds; stage 36, five toe buds; stages 37 and 38, lengthening of
toes. In all four stages, spiracle persistent, and pigmentation
resembling that of early stages.

_Stage 39._--Metatarsal tubercle present; greatest total length (36.9
mm.) attained.

_Stage 40._--Subarticular tubercles prominent; skin over forelimbs
transparent; cloacal tail-piece and spiracle absent; outer tooth-rows
degenerating; caudal fins shallower than in preceding stages; distal
part of tail nearly straight; size of dark markings on tail decreased;
pigment present on hind limb.

_Stage 43._--Forelimbs erupted; larval mouthparts absent; corner of
mouth between nostril and eye; transverse bands present on hind limbs;
tail greatly reduced (about 8 mm. in length).

_Stage 44._--Sacral hump barely noticeable; tail reduced to a stub;
corner of mouth at level of pupil of eye; dorsal surfaces pale
olive-green; venter white.

Changes proceed in a definite pattern during the growth and development
of tadpoles. Larval teeth are absent in hatchlings; the inner tooth-rows
develop first, and the third lower row last. At metamorphosis the third
lower row is the first to be lost. The tail increases gradually in
length relative to the body. In stage 25 the tail is 52.1 per cent of
the total length, and in stage 36, 64.6 per cent. In later stages the
tail becomes relatively shorter through resorption. Duellman and Klaas
(1964:320) noted a great size-variation in _Triprion_ tadpoles in stage
25. No such variation is apparent in any stage of any of the species of
_Smilisca_ studied.

The growth and development of the other species of _Smilisca_ do not
differ significantly from that of _S. phaeota_. The tadpoles of _S.
sila_ and _sordida_ from streams have relatively longer tails at
hatching. For example, in tadpoles of _S. sordida_ the average length of
tail is 64.0 per cent of the body-length in stage 25, and in stage 37,
67.0 per cent.

_Behavior_

The tadpoles of _S. baudini_, _cyanosticta_, _phaeota_, and _puma_ are
pelagic inhabitants of shallow ponds. Early stages of _S. baudini_ in
which external gills are present have been observed to hang vertically
with the gills spread out at the surface of the water, a behavior noted
by Zweifel (1964:206) in tadpoles of _Phrynohyas venulosa_, which also
develop in warm, standing water having a relatively low oxygen-tension.
When disturbed the pelagic tadpoles usually dive and seek shelter amidst
vegetation or in mud on the bottom. This behavior was observed in _S.
baudini_, _cyanosticta_, and _phaeota_ by day and at night. No tadpoles
of _S. puma_ were observed by day; those seen at night were near the
surface of small water-filled depressions in a grassy marsh; they
responded to light by taking refuge in the dense grass. Perhaps tadpoles
of this species are negatively phototactic and remain hidden by day.

The stream-inhabiting tadpoles of _S. sila_ and _sordida_ live in clear
pools in rocky streams, where they were observed to cling by their
mouths to rocks in the stream and to seek shelter amidst pebbles or
beneath rocks and leaves on the bottom. These tadpoles are not found in
shallow riffles.

We have not found tadpoles of two species of _Smilisca_ in the same body
of water and therefore cannot offer observations on ecological
relationships in sympatric situations.



PHYLOGENETIC RELATIONSHIPS


Identifiable hylid remains are known from the Miocene to the Recent, but
these fossils are mostly fragmentary and provide little useful
information regarding the phylogenetic relationships of living genera.
Frogs of the genus _Smilisca_ are generalized and show no striking
adaptations, either in their structure or in their modes of life
history.


Interspecific Relationships

In attempting to understand the relationships of the species of
_Smilisca_ we have emphasized osteological characters. The phylogeny
suggested by these characters is supported by other lines of evidence,
including external morphology, tadpoles, and breeding calls.

Our concept of the prototype of the genus _Smilisca_ is a moderate-sized
hylid having: (1) a well-developed frontoparietal fontanelle,
(2) frontoparietal lacking lateral processes, (3) no bony
squamosal-maxillary arch, (4) a fully ossified ethmoid, (5) paired
subgular vocal sac, (6) moderately webbed fingers and toes, (7)
relatively few supernumerary tubercles on the digits, (8) eggs deposited
in clumps in ponds, (9) anteroventral mouth in tadpoles bordered by one
row of labial papillae, but median part of upper lip bare, (10) tail
relatively short and deep in tadpoles, and (11) a breeding call
consisting of a series of like notes.

Two phyletic lines evolved from this prototype. The first of these was
the stock that gave rise to the _baudini_ group. The evolutionary
changes that took place in this line included increase in size,
development of a lateral curvature of the maxillary, and an increased
amount of cranial ossification, especially in the dermal roofing bones.
This phyletic line retained the larval characters and breeding call of
the prototype. The second phyletic line gave rise to the _sordida_ group
and diverged from the prototype in the development of an angular
maxillary and a breeding call consisting of a primary note followed by
secondary notes. The frogs in this phyletic line retained the moderate
size of the prototype and did not develop additional dermal bone. Our
concept of the phylogenetic relationships is shown graphically in Figure
17.

Within the _baudini_ group one stock retained separate nasals and did
not develop a bony squamosal-maxillary arch, but broad lateral processes
developed on the frontoparietals. The tadpoles remained unchanged from
the primitive type. This stock evolved into _S. phaeota_. In the other
stock the nasals became fully ossified and a bony squamosal-maxillary
arch developed. One branch of this second stock retained tadpoles having
only one row of labial papillae and did not develop lateral processes on
the frontoparietals; this branch evolved into _S. cyanosticta_. The
other branch diverged and gave rise to _S. baudini_ by developing
relatively shorter hind legs, large lateral processes on the
frontoparietals, and tadpoles having two rows of labial papillae.

Within the _sordida_ group the cranial features remained unchanged in
one line, which gave rise to _S. sila_, whereas in a second line the
nasals were reduced, and their long axes shifted with the result that
they are not parallel to the maxillaries; the amount of ossification of
the ethmoid was reduced, and the tadpoles developed two rows of labial
papillae. In this second line one branch retained the pond-breeding
habits and gave rise to _S. puma_, whereas a second branch became
adapted to stream-breeding and gave rise to _S. sordida_.

   _baudini_
      \   _cyanosticta_
       \ /
        +    _phaeota_       _sordida_
         \  /               /
           +        puma_  /
            \            \/   _sila_
             \           /_____/
              \        /
                \    /
                  \ /
                   |
                   |
                Prototype

    [Illustration: FIG. 17. Hypothesized phylogenetic relationships
       of the species of _Smilisca_.]

Certain aspects of this proposed phylogeny warrant further comment.
Features such as the deposition of additional bone that roofs the skull
or that forms lateral projections from the frontoparietals, like those
in _S. baudini_ and _phaeota_, are minor alterations of dermal elements
and not basic modifications of the architecture of the skull.
Consequently, we hypothesize the independent development of these dermal
changes in _S. baudini_ and _phaeota_. Similar kinds of dermal
modifications have evolved independently in many diverse groups of
frogs.

Likewise, we propose the parallel development of stream-adapted tadpoles
in _S. sordida_ and _sila_; in both cases the tadpoles adapted to
changing environmental conditions (see following section on evolutionary
history). Tadpoles of _S. sordida_ already had two rows of labial
papillae before entering the streams; subsequently the tadpoles
developed complete rows of papillae, ventral mouths and long tails
having low fins. Possibly the tadpoles of _S. sila_ had two rows of
labial papillae prior to their adapting to stream conditions; in the
process of adapting they developed ventral mouths and long tails having
low fins. Similar modifications in tadpoles have occurred in many
diverse groups of Middle American hylids, such as _Plectrohyla_,
_Ptychohyla_, the _Hyla uranochroa_ group, and the _Hyla taeniopus_
group.

Our lack of concern about coloration is due to the fact that, with the
exception of the blue spots on the flanks and posterior surfaces of the
thighs in some species, the coloration of _Smilisca_, consisting of a
pattern of irregular dark marks on a paler dorsum and dark transverse
bars on the limbs, is not much different from that of many other
Neotropical hylids. Blue is a structural color, rare among Amphibia,
which is achieved by the absence of lipophores above the guanophores.
Thus, the incident light rays at the blue end of the spectrum are
reflected by the guanophores without interference by an overlying yellow
lipophore screen. According to Noble (1931), lipophores are capable of
amoeboid movement that permits shifts in their positions, between or
beneath the guanophores. We do not know whether this behavior of
lipophores is widespread and is effected in response to environmental
changes, or whether it is a genetically controlled attribute that is
restricted in appearance. If the latter is the case we must assume that
the prototype of _Smilisca_ possessed such an attribute which was lost
in _S. baudini_, _phaeota_, and _puma_. The development of blue spots is
not constant in _S. sordida_ and _S. sila_; in _S. cyanosticta_ the
spots range in color from blue to pale green.

The coloration of the tadpoles is not distinctive, except for the
presence of dorsal blotches on the tails of _S. sila_ and _sordida_.
However, the similarity in pattern cannot be interpreted as indicating
close relationships because nearly identical patterns are present in
_Hyla legleri_ and some species of _Prostherapis_. This disruptive
coloration seems to be directly associated with the pebble-bottom,
stream-inhabiting tadpoles.

In the _baudini_ group, _S. phaeota_ and _cyanosticta_ are allopatric,
whereas _S. baudini_ occurs sympatrically with both of those species.
The call of _S. baudini_ differs notably from the calls of _S. phaeota_
and _cyanosticta_, which are more nearly alike. Although in the
phylogenetic scheme proposed here _S. sila_ is considered to be more
distantly related to _S. puma_ than is _S. sordida_, the calls of _S.
sila_ and _puma_ more closely resemble one another than either resembles
that of _S. sordida_. _Smilisca sila_ and _puma_ are allopatric, whereas
_S. sordida_ is broadly sympatric with both of those species. We assume
that in their respective phyletic lines the differentiation of both _S.
baudini_ and _sordida_ was the result of genetic changes in
geographically isolated populations. Subsequently, each species
dispersed into areas inhabited by other members of their respective
groups. Selection for differences in the breeding calls helped to
reinforce other differences in the populations and thereby aided in
maintaining specificity.


Evolutionary History

With respect to temporal and spatial aspects of evolution in _Smilisca_,
we have tried to correlate the phylogenetic evidence on _Smilisca_ with
the geologic data on Middle America presented by Lloyd (1963), Vinson
and Brineman (1963), Guzmán and Cserna (1963), Maldonado-Koerdell
(1964), and Whitmore and Stewart (1965). Likewise, we have borne in mind
the evidence for, and ideas about, the evolution of the Middle American
herpetofauna given by Dunn (1931b), Schmidt (1943), Stuart (1950, 1964)
Duellman (1958, MS), and Savage (MS).

According to Stuart's (1950) historical arrangement of the herpetofauna,
_Smilisca_ is a member of the Autochthonous Middle American Faunal
Element, and according to Savage's (MS) arrangement the genus belongs to
the Middle American Element, a fauna which was derived from a
generalized tropical American unit that was isolated in tropical North
America by the inundation of the Isthmian Link in early Tertiary, that
developed _in situ_ in tropical North America, and that was restricted
to Middle America by climatic change in the late Cenozoic.

Savage (MS) relied on the paleogeographic maps of Lloyd (1963) to
hypothesize the extent and centers of differentiation of the Middle
American Faunal Element. According to Lloyd's concept, Middle America in
the Miocene consisted of a broad peninsula extending southeastward to
about central Nicaragua, separated from the Panamanian Spur of
continental South America by shallow seas. A large island, the Talamanca
Range, and remnants of the Guanarivas Ridge formed an archipelago in the
shallow sea. The recent discovery of remains of mammals having definite
North American affinities in the Miocene of the Canal Zone (Whitmore and
Stewart, 1965) provides substantial evidence that at least a peninsula
was continuous southeastward from Nuclear Central America to the area of
the present Canal Zone in early mid-Miocene time. South America was
isolated from Central America by the Bolivar Trough until late
mid-Pliocene.

Thus, in the mid-Tertiary the broad peninsula of Nuclear Central
America, which consisted of low and moderately uplifted regions having a
tropical mesic climate, provided the site for the evolution of
_Smilisca_. It is not possible to determine when the genus evolved, but
to explain the differentiation of the species it is unnecessary to have
the ancestral _Smilisca_ present prior to the Miocene.

We view the Miocene _Smilisca_ as the prototype described in the
preceding section, and suppose that it lived in the mesic tropical
environment of the eastern part of the Central American Peninsula (in
what is now Costa Rica and western Panamá). Two stocks differentiated,
probably in middle Miocene times; one of these, the ancestral stock of
the _baudini_ group, was widespread on the Caribbean lowlands from the
Nicaraguan Depression to the Bolivar Trough, and the other, the
ancestral stock of the _sordida_ group, was restricted to the Pacific
lowlands of the same region. In late Miocene time the ancestral stock of
the _baudini_ group dispersed northwestward around the deep embayment in
the Nicaraguan depression into upper Central America (in what is now
Honduras and Guatemala) and thence into southern México. Apparently
differentiation took place on each side of the Nicaraguan Depression;
the frogs to the south of the depression evolved into _S. phaeota_,
whereas those to the north of the depression represented the stock from
which _S. baudini_ and _cyanosticta_ arose. Prior to the uplift of the
mountains in the late Miocene and the Pliocene the _baudini-cyanosticta_
stock probably was widespread in northwestern Central America. The
elevation of the mountains resulted in notable climatic changes,
principally the development of sub-humid environments on the Pacific
lowlands. The frogs living on the Pacific lowlands became adapted to
sub-humid conditions and developed into _S. baudini_. The stock on the
Caribbean lowlands remained in mesic environments and evolved into _S.
cyanosticta_.

Possibly in the middle Miocene before the Talamanca Range in Costa Rica
and western Panamá was greatly uplifted, the ancestral stock of the
_sordida_ group invaded the Caribbean lowlands of what is now Costa
Rica. The subsequent elevation of the Talamanca Range in the Pliocene
effectively isolated the ancestral stock of _S. sila_ on the Pacific
lowlands from the _puma-sordida_ stock on the Caribbean lowlands. The
former was subjected to the sub-humid conditions which developed on the
Pacific lowlands when the Talamanca Range was uplifted. It adapted to
the sub-humid environment by living along streams and evolving
stream-adapted tadpoles. On the Caribbean side of the Talamanca Range
the _puma-sordida_ stock inhabited mesic environments. The stock that
evolved into _S. puma_ remained in the lowlands as a pond-breeding frog,
whereas those frogs living on the slopes of the newly elevated mountains
became adapted for their montane existence by developing stream-adapted
tadpoles and thus differentiated into _S. sordida_.

Probably the six species of _Smilisca_ were in existence by the end of
the Pliocene; at that time a continuous land connection existed from
Central America to South America. The climatic fluctuations in the
Pleistocene, and the post-Wisconsin development of present climatic and
vegetational patterns in Middle America, brought about the present
patterns of distribution of the species. From its place of origin on the
Caribbean lowlands of lower Central America, _S. phaeota_ dispersed
northward into Nicaragua and southward along the Pacific slopes of
northwestern South America. Perhaps in the late Pleistocene or in
post-Wisconsin time when mesic conditions were more widespread than now,
_S. phaeota_ moved onto the Pacific lowlands of Costa Rica. Its route
could have been through the Arenal Depression. Subsequent aridity
restricted its range on the Pacific lowlands to the Golfo Dulce region.
Climatic fluctuation in northern Central America restricted the
distribution of _S. cyanosticta_ to mesic habitats on the slopes of the
Mexican and Guatemalan highlands and to certain humid areas on the
lowlands. _Smilisca baudini_ was well adapted to sub-humid conditions,
and the species dispersed northward to the Rio Grande Embayment and to
the edge of the Sonoran Desert and southward into Costa Rica. In
southern México and Central America the species invaded mesic habitats.
Consequently, in some areas it is sympatric with _S. cyanosticta_ and
_phaeota_.

_Smilisca puma_ dispersed northward onto the Caribbean lowlands of
southern Nicaragua. Its southward movements probably were limited by the
ridges of the Talamanca Range that extend to the Caribbean coast in the
area of Punta Cahuita in Costa Rica. _Smilisca sila_ dispersed along the
Pacific lowlands and slopes of the mountains from eastern Costa Rica and
western Panamá through eastern Panamá to northern Colombia. Climatic
fluctuation in the Pleistocene evidently provided sufficient altitudinal
shifts in environments in the Talamanca Range to permit _S. sordida_ to
move onto the Pacific slopes. From its upland distribution the species
followed streams down to both the Caribbean and Pacific lowlands, where
it is sympatric with _S. puma_ on the Caribbean lowlands and _S. sila_
on the Pacific lowlands.

The evolution of the species-groups of _Smilisca_ was effected through
isolation by physical barriers in the Cenozoic; the differentiation of
the species was initiated by further isolation of populations by changes
in physiography and climate. Present patterns of distribution resulted
from Pleistocene and post-Wisconsin climatic changes. Today, sympatric
species have different breeding habits and breeding calls which
reinforce the differences in morphology.



SUMMARY AND CONCLUSIONS


The genus _Smilisca_ is composed of six species of tree frogs; each
species is defined on the basis of adult morphology, larval characters,
and breeding behavior. Keys are provided to aid in the identification of
adults and of tadpoles.

Analysis of the characters and examination of type specimens indicates
that several currently-recognized taxa are synonymous, as follows:

  1. _Hyla beltrani_ Taylor, 1942 = _Smilisca baudini_.
  2. _Hyla gabbi_ Cope, 1876 = _Smilisca sordida_.
  3. _Hyla manisorum_ Taylor, 1954 = _Smilisca baudini_.
  4. _Hyla nigripes_ Cope, 1876 = _Smilisca sordida_.
  5. _Hyla wellmanorum_ Taylor, 1952 = _Smilisca puma_.

_Smilisca phaeota cyanosticta_ Smith, 1953 is elevated to specific rank,
and one new species, _Smilisca sila_, is named and described.

The skeletal system of developmental stages and the adult of _Smilisca
baudini_ is described, and the skull is compared with that of other
members of the genus.

The tadpoles are described, compared, and illustrated; the larval
development of _Smilisca phaeota_ is described.

Breeding behavior and breeding calls are described and compared. Some
species of _Smilisca_ have breeding choruses. Two species, _S. sila_ and
_sordida_, breed in streams, whereas the others breed in ponds.

The genus is considered to be part of the Middle American Faunal
Element; the species are thought to have differentiated in response to
ecological diversity and historical opportunities provided by Cenozoic
changes in physiography and climate.



LITERATURE CITED


BAIRD, S. F.

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BALDAUF, R. J.

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BARBOUR, T.

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BLAIR, W. F.

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BOGERT, C. M.

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BRATTSTROM, B. H.

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BREDER, C. M., Jr.

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BROCCHI, P.

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CARVALHO, A. L.

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COCHRAN, D. M.

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COPLAND, S. J.

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DUELLMAN, W. E.

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DUELLMAN, W. E. and KLAAS, L. T.

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DUMÉRIL, A. M. C. and BIBRON, G.

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DUNN, E. R.

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FOUQUETTE, M. J., JR.

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FUNKHOUSER, ANNE

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GADOW, H.

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GAIGE, H. T., HARTWEG, N. and STUART, L. C.

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GOSNER, K. L.

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GRIFFITHS, I.

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GUZMÁN, E. J. and CSERNA, Z.

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HECHT, M. K.

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JOHNSON, C.

  1959.  Genetic incompatibility in the call races of _Hyla versicolor_
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MALDONADO-KOERDELL, M.

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MASLIN, T. P.

  1963.  Notes on some anuran tadpoles from Yucatán, México. Herpetologica,
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MITTLEMAN, M. B. and LIST, J. C.

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_Transmitted March 14, 1966._



  []
  31-3430



UNIVERSITY OF KANSAS PUBLICATIONS

MUSEUM OF NATURAL HISTORY


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               $1.00.

           16. A new subspecies of the fruit-eating bat, Sturnira
               ludovici, from western Mexico. By J. Knox Jones, Jr.,
               and Gary L. Phillips. Pp. 475-481, 1 figure in text.
               March 2, 1964.

           17. Records of the fossil mammal Sinclairella, Family
               Apatemyidae, from the Chadronian and Orellan. By William
               A. Clemens, Jr. Pp. 483-491, 2 figures in text.
               March 2, 1964.

           18. The mammals of Wyoming. By Charles A. Long. Pp. 493-758,
               82 figures in text. July 6, 1965. $3.00.

           Index. Pp. 759-784.


  Vol. 15.  1. The amphibians and reptiles of Michoacán, México. By
               William E. Duellman. Pp. 1-148, pls. 1-6, 11 figures in
               text. December 20, 1961. $1.50.

            2. Some reptiles and amphibians from Korea. By Robert G.
               Webb, J. Knox Jones, Jr., and George W. Byers.
               Pp. 149-173. January 31, 1962.

            3. A new species of frog (genus Tomodactylus) from western
               México. By Robert G. Webb. Pp. 175-181, 1 figure in text.
               March 7, 1962.

            4. Type specimens of amphibians and reptiles in the Museum
               of Natural History, The University of Kansas. By William
               E. Duellman and Barbara Berg. Pp. 183-204.
               October 26, 1962.

            5. Amphibians and reptiles of the rainforests of southern
               El Petén, Guatemala. By William E. Duellman. Pp. 205-249,
               pls. 7-10, 6 figures in text. October 4, 1963.

            6. A revision of snakes of the genus Conophis (Family
               Colubridae, from Middle America). By John Wellman.
               Pp. 251-295, 9 figures in text. October 4, 1963.

            7. A review of the Middle American tree frogs of the genus
               Ptychohyla. By William E. Duellman. Pp. 297-349,
               pls. 11-18, 7 figures in text. October 18, 1963.
               50 cents.

            8. Natural history of the racer Coluber constrictor.
               By Henry S. Fitch. Pp. 351-468, pls. 19-22, 20 figures
               in text. December 30, 1963. $1.00.

            9. A review of the frogs of the Hyla bistincta group.
               By William E. Duellman. Pp. 469-491, 4 figures in text.
               March 2, 1964.

           10. An ecological study of the garter snake, Thamnophis
               sirtalis. By Henry S. Fitch. Pp. 493-564, pls. 23-25,
               14 figures in text. May 17, 1965.

           11. Breeding cycle in the ground skink, Lygosoma laterale.
               By Henry S. Fitch and Harry W. Greene. Pp. 565-575,
               3 figures in text. May 17, 1965.

           12. Amphibians and reptiles from the Yucatan Peninsula,
               México. By William E. Duellman. Pp. 577-614, 1 figure
               in text. June 22, 1965.

           13. A new species of turtle, genus Kinosternon, from Central
               America. By John M. Legler. Pp. 615-625, pls. 26-28,
               2 figures in text. June 20, 1965.

           14. A biogeographic account of the herpetofauna of Michoacán,
               México. By William E. Duellman. Pp. 627-709, pls. 29-36,
               5 figures in text. December 30, 1965.

           15. Amphibians and reptiles of Mesa Verde National Park,
               Colorado. By Charles L. Douglas. Pp. 711-744, pls. 37
               and 38, 6 figures in text. March 7, 1966.

           Index in preparation.


  Vol. 16.  1. Distribution and taxonomy of mammals of Nebraska. By J.
               Knox Jones, Jr. Pp. 1-356, plates 1-4, 82 figures in
               text. October 1, 1964. $3.50.

            2. Synopsis of the lagomorphs and rodents of Korea. By J.
               Knox Jones, Jr., and David H. Johnson. Pp. 357-407.
               February 12, 1965.

            3. Mammals from Isla Cozumel, Mexico, with description of
               a new species of harvest mouse. By J. Knox Jones, Jr.
               and Timothy E. Lawlor. Pp. 409-419, 1 figure in text.
               April 13, 1965.

            4. The Yucatan deer mouse, Peromyscus yucatanicus.
               By Timothy E. Lawlor. Pp. 421-438, 2 figures in text.
               July 20, 1965.

            5. Bats from Gautemala. By J. Knox Jones, Jr. Pp. 439-472.
               April 18, 1966.

           More numbers will appear in volume 16.


  Vol. 17.  1. Localities of fossil vertebrates obtained from the
               Niobrara Formation (Cretaceous) of Kansas. By David
               Bardack. Pp. 1-14. January 22, 1965.

            2. Chorda tympani branch of the facial nerve in the middle
               ear of tetrapods. By Richard C. Fox. Pp. 15-21.
               June 22, 1965.

            3. Fishes of the Kansas River System in relation to
               zoogeography of the Great Plains. By Artie L. Metcalf.
               Pp. 23-189, 4 figures in text, 51 maps. March 24, 1966.

            4. Factors affecting growth and production of channel
               catfish, Ictalurus punctatus. By Bill A. Simco and
               Frank B. Cross. Pp. 191-256, 13 figures in text.
               June 6, 1966.

            5. A new species of fringe-limbed tree frog, genus Hyla,
               from Darién, Panamá. By William E. Duellman. Pp. 257-262,
               1 figure in text. June 17, 1966.

            6. Taxonomic notes on some Mexican and Central American
               hylid frogs. By William E. Duellman. Pp. 263-279.
               June 17, 1966.

            7. Neotropical hylid frogs, genus Smilisca. By William E.
               Duellman and Linda Trueb. Pp. 281-375, pls. 1-12,
               17 figures in text. July 14, 1966.

           More numbers will appear in volume 17.



Transcriber's Note

With the exception of the corrections listed below and several minor
corrections not listed, the text presented is that which appeared in the
original printed version. The list of Kansas University publications has
been compiled at the end of the article.


Typographical Corrections

  Page    Correction
  ====  ==================
   287  cleared => cleaned
   292  Data based of => Data based on
   298  CNMH => CNHM
   299  Acahuitzotla => Acahuizotla
   304  cyanostica => cyanosticta
   305  Quatemala => Guatemala
   307  cyanostica => cyanosticta
   313  Matagalapa => Matagalpa
   322  Carribean => Caribbean
   323  Centralia => Centrali
   336  proportionaely => proportionately
   346  noticably => noticeably
   362  Fouquett => Fouquette
   372  CARVALJO => CARVALHO
   375  Dumeril => Duméril
    ii  trutles => turtles





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