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SMITHSONIAN
INSTITUTION
MUSEUM
O F

NATURAL
HISTORY

Brother Niceforo Maria, director of Museo del Instituto de La

whom

this

book

is

dedicated.

Salle,

Bogota, to

UNITED STATES NATIONAL MUSEUM BULLETIN

Frogs of Colombia

DORIS M. COCHRAN
Curator, Division of Reptiles and Amphibians,

United States National

Museum

and

COLEMAN

J.

GOIN

Department of Biological Sciences, University of Florida,

and Research Associate, Carnegie Museum

SMITHSONIAN INSTITUTION PRESS
CITY OF WASHINGTON
1970

288

Publications of the United States National

The
series,

Museum

scientific publications of the United States National Museum include two
Proceedings of the United States National Museum and United States National

Museum

Bulletin.

In these series are published original articles and monographs dealing with
the collections and work of the Museum and setting forth newly acquired facts
in the fields of anthropology, biology, geology, history, and technology. Copies
of each publication are distributed to libraries and scientific organizations and
to specialists and others interested in the various subjects.
The Proceedings, begun in 1878, are intended for the publication, in separate
form, of shorter papers. These are gathered in volumes, octavo in size, with
the publication date of each paper recorded in the table of contents of the volume.
In the Bulletin series, the first of which was issued in 1875, appear longer,
separate publications consisting of monographs (occasionally in several parts)

and volumes

which are collected works on related subjects. Bulletins are
on the needs of the presentation.
Since 1902, papers relating to the botanical collections of the Museum have been
published in the Bulletin series under the heading Contributions from the United
in

either octavo or quarto in size, depending

States National Herbarium.

This work forms number 288 of the Bulletin

series.

Frank

A.

Taylor

Director, United States National

Museum

UNITED STATES GOVERNMENT PRINTING OFFICE
WASHINGTON
1970
:

For

sale

by the Superintendent

of

Documents, U.S. Government Printing

Washington, D.C. 20402

-

Price $4

Office

Dedicated

to

Brother

Niceforo

Maria,

whose

pioneering work in Colombian herpetology has
cleared a path that
follow.

makes

it

easier for those

who

Contents
Page

Introduction
Class Amphibia, Superorder Salientia, Order Anura
Suborder Aglossa
Family Pipidae
Pipa Laurenti
P. parva Ruthven and Gaige
P. snethlageae Midler
P. pipa (Linnaeus)
Suborder Diplasiocoela

1

7
7

7
7

8
9

10
12
13
13
13
14
16

Family Ranidae
Subfamily Dendrobatinae
Dendrobates Wagler
D. Irivittatus (Spix)
D.
D.
D.
D.
D.
D.
D.
D.
D.
D.

ingeri,

new

species

hahneli Boulenger

17

Schmidt
opisthomelas Boulenger
minutus ventrimaculatus Shreve
minutus minutus Shreve
tinctorius wittei Laurent
lugubris O.

19

22
23
25
25
28
31
32
34
35
38
40
42

Berthold
(Posada Arango)

tinctorius histrionicus
tinctorius chocoensis

Funkhouser
and Bibron

tinctorius confluens

Phyllobates Dumeril

P. bicolor Bibron
P. boulengeri (Barbour)

P.
P.
P.
P.
P.
P.
P.
P.
P.
P.
P.
P.

femoralis (Boulenger)

mertensi Cochran
thorntoni,

new

and Goin

44
46
48
52

species

chocoensis (Boulenger)

brunneus (Cope)
palmatus Werner
pratti Boulenger
latinasus

55
57
59
62
65
68
70
70
70
74
74
74

Cope

inguinalis (Cope)

subpunctatus subpunclatus (Cope)
subpunctatus walesi, new subspecies
vergeli

(Hellmich)

Subfamily Raninae
Rana Linnaeus
R. palmipes Spix

Family Microhylidae
Relictivomer Carvalho
R. pearsei (Ruthven)
VII

B. A. A. B. — Continued 119 120 122 123 125 127 128 130 131 133 135 137 139 140 141 144 145 146 146 149 . aterrimum Giinther Clenophryne Mocquard C. A. ovalis (Schneider) Glossosloma Giinther G. Superorder Salientia. B. new subspecies granulosus beebei Gallardo granulosus humboldti Gallardo typhonius (Linnaeus) marinus (Linnaeus) blombergi coniferus Myers and Funkhouser Cope sternosignatus Giinther Cope subspecies ignescens (Cornalia) varius subornaius Werner pedimarmoratus Rivero nicefori Rivero spurrelli Boulenger varius elegans (Boulenger) longibrachius Rivero varius bibroni (O. B. A. paradoxa nicefori. A. geayi 80 80 82 83 85 85 Mocquard Otophryne Boulenger 0. ceraiophrys Boulenger B. A. B. A. A. A. B. A. A. robusta Boulenger 88 Suborder Procoela Family Pseudidae Pseudis Wagler new P. 92 93 95 97 98 101 105 107 113 114 116 glaberrimus Giinther haematiticus hypomelas Boulenger Family Atelopodidae Atelopus Dum6ril and Bibron A. A. 88 89 89 91 B. A. Order Suborder Diplasiocoela Family Microhylidae Elachistocleis — Continued — Continued Anura Page Parker 77 77 E. Schmidt) carrikeri Ruthven varius glyphus longirostris Dunn Cope walkeri Rivero Family Hylidae Agalychnis Cope A. pachydermus (O.CONTENTS VIII Class Amphibia. B. new calcarifer spurrelli Boulenger Boulenger 109 Ill 118 118 bufoniformis Peracca ebenoides ebenoides River o ebenoides marinkellei. A. B. B. Schmidt) A. B. B. A. subspecies Family Bufonidae Bufo Laurenti 91 rostratus rostratus Noble rostratus nicefori.

columbiana Boettger H. rubracyla. Superorder Salientia. H. monticola argenteovirens (Boettger) Hyla H. bogerti. guibei. new species H. new species species G.CONTENTS Class Amphibia. H. H. IX species H. new G. new 151 151 153 155 156 158 161 163 164 Gastroiheca Fitzinger G. albomarginata Spix H. new species H. rubra Daudin H. labialis labialis Peters H. rubeola. bufona. new species H. johnsoni Ruthven new species Cryptobalrachus C. Lutz H. punctata (Schneider) H. variabilis Boulenger H. labialis platydactijla Boulenger H. C- fuhrmanni (Peracca) C. Order Suborder Procoela Continued — Family Hylidae— Continued Anura Noble C. mertensi. H. new species H. nicefori Gaige G. boulengeri Ruthven new species G. albopunctulata Boulenger H. Laurenti lanciformis (Cope) crepitans Wied rosenbergi Boulenger maxima (Laurenti) geographica geographica Spix H. aureomaculala. phantasmagoria Dunn H. buckleyi Boulenger H. boulengeri (Cope) H. new species H. parviceps Boulenger 167 170 172 175 177 181 185 188 189 192 196 199 203 206 209 210 213 215 218 220 222 225 226 228 229 231 233 236 239 243 245 248 251 254 256 258 261 264 266 270 . granosa Boulenger H. cabrerai. palpebrogranulata Andersson H. cornutum (Boulenger) G. new species G. helenae Dunn G. nicefori. medemi. squalirostris A. carri. foliamorta Fouquette H. calcarata Troschel H. rhodoporus Giinther Hyla sp H. — Continued p age Jimmez de la Espada proboscidea Jimmez de la Espada Cerathyla C. labialis krausi Hellmich H. bokermanni Goin H.

sarayacuensis Shreve H. leprieuri (Dum6ril and Bibron) Phrynohyas Fitzinger P. ingens Duellman P. orccsi . new species H.CONTENTS X Class Amphibia. rossalleni Goin H. minuta Peters H. aurantiacus (Daudin) S. laynei Goin H. Superorder Salientia. P. reticulata Jimfnez de la Espada H. P. gabbi (Cope) Sphaenorhynchus Tschudi S. bicolor (Boddaert) P. vcnulosa (Laurenti) Phyllomedusa Wagler P. mathiassoni. new species H. P. jordani Stejneger and Test Family Leptodactylidae Subfamily Leptodactylinae Amblyphrynus Cochran and Goin A. 272 274 277 280 284 286 290 295 296 299 302 304 306 308 311 Barbour boliviana Boulenger tarsius (Cope) P. marmorata (Laurenti) H. leucophyllata Beireis H. riveroi. dorisae (Goin) Tetraprion Stejneger and Test T. hobbsi. membranacea Andersson H. edentula Andersson Phyllomedusa sp Smilisca Cope S. P. new species H. phlebodes Stejneger H. phaeota (Cope) S. bogolensis (Peters) Osleocephalus Steindachner Funkhouser O. ebraccata Cope H. taurinus Steindachner O. ingeri Cochran and Goin Borborocoetes Bell B. new species H. habrus (Goin) S. microcephala Cope H. Order Suborder Procoela Continued — Family Hylidae — Continued Anura — Continued Page H. columbianus Werner 317 317 319 322 324 324 326 330 330 332 334 337 338 340 342 344 345 346 348 350 351 353 355 358 358 359 360 360 360 363 363 . blombergi Funkhouser hypochondrialis hypochondrialis (Daudin) nicefori • • 313 313 Hyloscirtus Peters O. oliveae.

Superorder Salicntia. w-nigrum (Boettger) E. lehmanvalenciae Thornton E. Order — XI — Continued Anura Suborder Procoela Continued Family Leptodactylidae Continued Ceratophrys — Wied C. calcaratus (Boulenger) E. cornuta (Linnaeus) Edalorhina Jiminez de la Espada E. vertebralis (Boulenger) E. rhodomystax (Boulenger) L. nicefori. elegans (Peters) E. affinis (Werner) E. anomalus (Boulenger) E. cornutus cornutus (Jimfnez de la Espada) Eupemphix Steindachner E. insignitus Ruthven E. delicatus Ruthven E. new subspecies E. bogotensis (Peters) E.CONTENTS Class Amphibia. pentadactylus (Laurenti) L. surdus cabrerai. lymani Barbour and Noble E. schmidti (Cochran and Goin) Leptodadylus Fitzinger L. latidiscus tamsitti. diastema (Cope) E. pustulosus (Cope) Hydrolaetare Gallardo H. taeniatus (Boulenger) E. gularis (Boulenger) E. terraebolivaris Rivero E. bufoniformis (Boulenger) E. longirostris (Boulenger) E. brederi Dunn E. calcarata Boulenger C. Ichmanni (Boettger) E. mystaceus (Spix) Page 364 364 366 368 369 371 375 376 380 382 384 387 389 391 393 395 398 399 401 403 405 407 409 411 413 416 418 421 423 424 426 428 430 433 435 437 439 441 442 444 444 448 448 449 450 452 455 457 459 . new species E. new subspecies E. insularum Barbour L. palmeri (Boulenger) E. erythropleurus (Boulenger) E. frater (Werner) E. carmelitae Ruthven E. conspicillatus (Giinther) E. raniformis (Boulenger) E. cruentus (Peters) E. sandae-martae Ruthven E. latidiscus latidiscus (Boulenger) E. megalops Ruthven E. perezi Jimfnez de la Espada Eleutherodactylus Dume>il and Bibron E. marmoratus hylaedadylus (Cope) L.

geckoideum Jimfnez de la Noble C. poecilochilus diptychus Boulenger L. brachyops (Cope) P. lineatus (Schneider) Niceforonia Goin and Cochran N.XH CONTENTS Class Amphibia. poecilochilus poecilochilus (Cope) L. new subspecies Pleurodema Tschudi P. medemi new species new species new species C. podicipinus peter si (Steindachner) Lithodytes Fitzinger L. walkeri Ruthven Family Centrolenidae Centrolene Jimfnez de la Espada C. myersi Goin and Cochran Subfamily Rhinodermatinae Geobatrachus Ruthven G. Collection localities Species Distribution References Espada . Order — Anura Suborder Procoela Continued Family Leptodactylidae Continued — — Continued Page Plates 462 464 466 468 471 473 475 475 478 478 480 480 483 483 486 488 489 491 491 493 493 494 496 496 496 499 500 502 506 508 511 513 515 517 537 545 573 Index 641 L. buckleyi (Boulenger) C. johnelsi. nana Goin and Cochran Physalaemus Fitzinger P. fleischmanni (Boettger) C. C. neglectus dunni. intermedins A. sagittifer (O. pusilla (Ruthven) Trachyphrynus Goin and Cochran T. parabambae (Boulenger) C. antioquiensis Noble Centrolenella C. sibilatrix (Weid) L. Lutz L. Superorder Salientia. Schmidt) Pseudopaludicola Miranda-Ribeiro P. grandisonae. rubido Cope L.

very best wishes for your personal welfare and believe me. We worked closely throughout the project. The specimens were collected chiefly along the Amazonian River (Leticia). and Lepto- dactylidae were largely prepared by the senior author. D. my offer and would thank you for a prompt answer. Ranidae. Colombia Dr. it was necessary that we divide the work Of the accounts that to follow. and Centrolenidae by the junior author. and a tag or label bearing a number would be attached to each of the specimens I and corresponding to I wonder my own list. Florida. The land area of Colombia.136.C. you can accept if Please accept my Yours very sincerely. Bufonidae. D.Introduction This Bulletin had was letter origin its on July 9. comprising 1.C. Hylidae. Museum on the we soon found collec- that there were rather extensive collections available in other institutions and seemed advisable to bring all of the material together and present a single. thus. 1957. With one of us working in Washington. we are jointly responsible for all parts. those of the Microhylidae. Villavicencio. especially frogs and toads. to 4°14' S.. the ones of the families Pipidae. Caqueta.D. Cochran United States National Washington. comprehensive report. to 79°01' W. and the other in it Gainesville. and from longitude 66°51' W. extends from latitude 12°30' N. the Cucuta region in Norte de Santander.E. etc. however. The material sent for study would be left as a gift for the National Museum. Brother NiceToro Maria Director of the Although tion in the it was our Museo original intention to report only del Instituto de La Salle. Atelopodidae. some extent. Doris M. Museum Madam: have before me your important publication "Frogs of Southeastern Brazil" this prompts me to ask you whether you could accept to identify my collection of Batrachians. received at the United States National Museo when the Museum: following del Institute* de La Salle Bogota . Meta. and near Floiencia.153 square kilometers. and we families tried to solve all "problems" together. . Pseudidae.

704) Norte de Santander (20. and Cerro Basin. The Cordillera Oriental extends northward in Colombia as the Sierra de Motilones. NATIONAL MUSEUM BULLETIN 88 2 Several geographic features of Colombia seeni to contribute to its immense frog fauna. Reaching as it does from the Amazon River to the Caribbean Sea and from the Orinoco River to the Pacific Ocean. and the Cordillera Occidental.317) (86.S. Its northern shores touch on the Caribbean.201) Valle (21.043) Narino (32. South of latitude 9°05' North.736) Cauca (30. The western half of Colombia is very rugged.370) Atlantico (3.800 meters. and Montes de Oca. The Andean Chain Ecuador as a single range but soon divides enters the country from into three ranges: the Cordillera Oriental.750 meters in the Cordillera Central. In Colombia's southern Andes (before they divide) the Nevada de Cumbal attains a height of 4. the Serrania de Valledupar.094) Huila (19.444) Choc6 (46.632) Boyaca" (67." and four "comisarias. which. in the Orinoco Basin east of the Andes in Meta. the Departa- mento Norte de Santander lies east of these ranges in the Maracaibo The highest peak in Colombia is the Chundua or Pico Crist6bal Coldn.080) Cundinamarca (23.493 meters in the Cordillera Oriental.541) Santander (30.2 U. and the only one that borders upon both oceans. Tamana 4. which reaches an altitude of 5.742) Guajira (21. Smaller mountain massifs are the equally isolated Serrania de La Marcarena. form the political divide between Colombia and Venezuela.669) Intendencias Arauca (22. its western on the Pacific. which during part of the 19th century was known as New Granada.866) Tolima (23.928) Caldas (13. it derives its fauna from many sources. In the northwest is the isolated Sierra Nevada de Santa Marta.020) C6rdoba (23. In the three Andean chains. Colombia.869) Caqueta (96. the Nevado del Huila reaches 5. which separates the Rio Atrato drainage from the Pacific Ocean in northern Choc6 and is more or less contiguous with the Serrania del Darien of Panama." five "intend encias.786) Meta Magdalena San Andres y Providencia (46.288) BoHvar (37. and the Serrania de Baud6.200 meters in the Cordillera Occidental. the Cordillera Central. in the Sierra Nevada de Santa Marta.480) (43) . north of latitude 9°05' North." These (with their respective areas in square kilometers) are listed below: Depaktamentos Antioquia (64. It is the only South American country directly connected to Central America through the Isthmus of Panama. the Nevado de Ritacuva 5. is now divided politically into sixteen "departamentos.764 meters.

the junior author presents a diagnosis before each description.. Dr. Ross Allen. Robert F. The senior author omits a diagnosis before discussing critical length. from base of thumb to tip of third ringer. Jean Guibe\ Museum National d'Histoire Naturelle. Academy of Natural Sciences of Philadelphia. Paris. Also. Charles M. M. the junior author measures in tenths of millimeters. Rijksmuseum van Natuurlijke Historie. Richard Zweifel. one-half the distance between distal ends of femur when at right angles to body. Alf G. Duellman. Boeseman. Dr. Alice G. Gottingen. head width. Naturhistoriska Riksmuseum. Dr. Florida State Museum. Zoologisches Institute. tibia. University of Florida. The measurements are the head-and-body length. Zoologische Staatssammlung. University of Kansas. Dr. Carlos Lehmann. femur. although the value of such precise naming for old and faded material is often questionable. Eiselt. Dr. Bogota. 1886 edition. Stockholm. American Museum of Natural History. Jose Gallardo. Vienna. Inger and Hymen Marx. previously described forms. Leviton. Sao Paulo. We are deeply indebted to the following for the privilege of examining specimens in their care and for other courtesies: E. Johnels. J. Naturhistorisches Museum. Bogert and Dr. P. Antenor Carvalho. Museo de Historia Natural. Bertha Lutz and Dr. head from posterior corner of tympanum..217) (18. and hand.061) For color descriptions the senior author has used the color names Robert Ridgway's A Nomenclature of Colors for Naturalists . from distal base of inner metatarsal tubercle to tip of fourth toe. Goteborg. C. Sven Mathiasson. popular color names.784) Vichada (98. Naturhistoriska Museet. Chicago Natural History Museum. Dr.238) (120. Werner C. Dr. California Academy of Sciences. C. Grandison and J. Dr. Dr. Our pleasure in acknowledging the help we have received while working on this project is tempered only by the fear that we have failed to thank some of the many persons who have been so kind to us. . Robert Mertens and Dr. Munich. Battersby. British Museum (Natural History). Museo Argentina de Ciencias Naturales. Dr. Dr. Popay&n. Rio de Janeiro. Dr.FROGS OF COLOMBIA — COCHRAN AND GOIN 3 COMISABIAS Amazonas Putumayo Vaup&s (167. The senior author records measurements to the nearest half millimeter. Walter Hellmich. Universidad de Los Andes. Dr. Museu National. Ross Allen's Reptile Institute. . Kuenzer. James Bohlke and Edmond Malnate. Walter Auffenberg and Dr. The junior author uses more in . between typani. Natur-Museum. A. William E. Alice Hunter. considering most diagnostic characters to have appeared in the key to the genus. William Riemer. Bokermann. foot. Alan E. Konrad Klemmer. Dr.

Goin. Secretaria da Agricultura. Dr. and Mr. Niceforo Maria. Juan A. 1 The manuscript was in the hands of the editor at the time Dr. Dr. Hno.* formerly of San Fernando State College and now of the United States National Museum. we are deeply grateful to the National Science Foundation for the following grants: Doris M. Cambridge. We cannot thank her enough. Dr. M. University of Kansas. Dr. The National Science Foundation made it possible for us to visit museums both in this country and abroad. To our numerous friends in Colombia. Cali. Rivero. and Coleman J.. George Zug. whose kindness not only made our visits profitable but made them exceedingly pleasant and enlightening. Walker. and gave us an opportunity to visit and collect in Colombia. Quito. C. University of Southern California. Departamento de Zoologia. Jean Vellard. University of Florida. Thornton. Stanford University Natural History Museum. and we are grateful to him. Goin spent weeks poring over maps to trace down obscure localities and she also typed much of the earlier drafts of the manuscript. G-5628. Dr. Universidad de Los Andes. Gerardo Reichel-Dolmatoff of Bogota made for us several significant collections for which we are most grateful. Carnegie Museum. Charles F. Dr. Olive B. Federico Medem made special efforts to collect for us in out-ofthe-way places on numerous occasions. contributed materially to this study on more than one occasion. Dr. Jay Savage. and for these carefully executed figures the artists have our thanks. Peters." Lima. Dr. Dr.G illness. For this we cannot thank him enough. Williams. Mr. Taylor. University of Puerto Rico at Mayaguez. Wilmot A. — . James A. nuestros mas sinceros agradecimentos. James A. Museum of Zoology. University of Michigan. artists in the Department of Biology. University of Michigan. Bogota. Universidad del Valle. Universidad Central de Venezuela. Dr. Dr.. Sao Paulo. Richmond. Tamsitt. Museum of Comparative Zoology. E. often at his own expense. and GB-1339. Museo de Historia Natural "Javier Prado. Dr. Escuela Politecnica Nacional.J. graduate assistant at the University of Florida. Gustavo Orces V. Paid Laessle. Dr. Cochran. Museo del Instituto de La Salle. I cannot thank him enough for the diligent and careful labor he contributed to the final preparation of this work. Most of the drawings contained herein are the works of Miss Ester Coogle and Mr. Paulo Vanzolini. Myers. GB-407 (201D-294). George S. Thus. Dr. Peters at the United States National Museum. Ernest E. Bogota. the chore of checking the thousand and one details necessary in a job of this magnitude fell on the shoulders of Dr. H. G-8625. NATIONAL MUSEUM BULLETIN 288 Senckenberg. Dr. Museum of Zoology. Cochran came down with her terminal and since I was in Gainesville. Crawford Jackson. Graham Netting and Neil D. Museo de Biologia. Dr.S. G-13325. Caracas.4 U. Juan Racenis.

Surely there are a number of undescribed species remaining to be discovered in Colombia. Quito Collection of Federico Medem. Chicago Departmento de Zoologia. they are not known 337-262—70 2 to occur there at the present time. J. University of Kansas. Universidad del Cauca. Washington UV Universidad del Valle. Palo Alto NHMW Naturforschende Gesellschaft]. A. in the Universidad de los Andes. ZIMUG Zoologisches Institut ZSM Zoologische Staatssammlung. Vienna Naturhistoriska Museet. Brazil und Museum der Universitat. Bokermann. Sccretaria da Agricultura. London of Sciences. Neill Werner C. Pittsburgh Chicago Natural History Museum. of New York City (Natural History). Stockholm Rijkemuseum van Natuurlijke Historie. "Javier Prado. University of Michigan. Tamsitt Ciencias Naturales. Universidad Central de Venezuela USNM University of Florida. Goin Bogotd CNHM DZSP EK EPN Collection of Eugene Kramer Escuela Politecnica National. Bogotd MN MNHUK MZUM National." of Museum Lima James R. Ann Arbor NM NMS Naturhistorisches Museum. Sao Paulo Academy California Coleman J. Peters. Gottingen Munich Although we here recognize 212 species and subspecies of Colomwe are not deluded into believing that our list is final in any way. . Gainesville University of Puerto Rico. in FM the United States National Museum IOC JAP Instituto JP Museo de JRT Collection of MACN Museo Argentina de MCZ Museum Museum Oswaldo Cruz.FROGS OF COLOMBIA The —COCHRAN AND GOLN 5 following abbreviations are used to identify the institutions and individuals owning or having custody of the specimens examined: AMNH American Museum ANSP Academy of Natural Sciences of Philadelphia BM British Museum Natural History. Popayan Biologfa. Mayagiiez United States National Museum. Rio de Janeiro James A. Leiden Stanford University Natural History Museum. Rio de Janeiro Museum of Natural History. Buenos Aires Comparative Zoology. Marinkelle. Cambridge National d'Histoire Naturelle. in the United States National Collection of MHNP MLS Museo Museu Historia Natural. Cali WCAB Collection of WTN Collection of Wilfred T. UC UCV UF UPR Museo de Museo de Historia Natural. and probably there are dozens of known species that someday will be recorded from that country although bian frogs. CM Carnegie Museum. San Francisco CAS CJG Collection of CJM Collection of C. Lawrence Museum of Zoology. in Silo Paulo. Paris del Instituto de La Salle. Goteborg Natur-Museum Senckenberg [Senckenbergische NR RNH SUNHM Frankfurt-am-Main Naturhistoriska Riksmuseum.

S. NATIONAL MUSEUM BULLETIN 2 88 visualized our task not as one of preparing the frogs of Colombia but rather now known and making such knowledge If this report makes the study of as bringing together available in a single Colombian frogs easier in has been in the past. then this labor of love without value and no further apology is necessary. the future than it is not .6 U. final what is volume. we have word on the Thus.

toes completely webbed. subcylindrical and rounded. no lateral flaps at angle of mouth. of the toes. 1768. but fused to vertebrae in adult. Tynpanum lacking. no horny tips on bl any mouth present. Sternum Eggs develop in pits in the dorsal skin of the female. tentacle-like skin structures near mouth and on throat. P. Genus Pipa Laurenti Pipa Laurenti. Dia- dilated pophyses of sacral vertebra strongly dilated. many short. Coracoids and precoracoids strongly divergent. Fingers free. Omosternum rudimentary.. lateral flaps at angle of heel reaching to axilla. not overlapping. Key a No 1 to Colombian Species of Pipa inner metatarsal tubercle. but no large flap-like appendages at tip of P. the former much and connected by a broad double cartilage. size to between eye and 45. brown horny tips on the first three toes. heel reaching tip of snout. americana Laurenti =Rana pipa Linnaeus) — Pupil round. Frogs of Colombia Class Superorder AMPHIBIA SALIENTIA Suborder : Order ANURA AGLOSSA No true tongue present. snethlageae snout and angle of jaw. Family Pipidae Tips of fingers divided to form star-like sensory structures. size to 88 Body mm 7 . Terminal phalanges pointed. or articulated to it by a single condyle. 24 (type species. p. Palate smooth. Vertebrae opisthocoelous. Sternum a large cartilaginous plate.5 An a2 mm P. Ribs free in tadpole. Outer metatarsals separated. Generic diagnosis. Shoulder girdle partly or wholly firmisternal. parva inner metatarsal tubercle present. small. Urostyle fused to sacrum.

in postaxillary region more than 1% times the greatest width of head. heel reaches anterior insertion of shoulder. 3. Toes long and slender. tapering. 1937b. Body very stout.— Gorham. When hind leg is adpressed. 3. 9. interorbital diameter about 1% times the snout length. 1939b. their order of length being 1. 1937. 1 (type locality. . terminal. 5. 1963. the tubercles becoming elongate and tipped with small spines on the limbs. when limbs are laid along the sides. pipa Pipa parva Ruthven and Gaige Plate 1a-c 1923. rounded when viewed from above. p. — USNM Description. 41.— Gin^s. Skin of upper parts with small.— Dunn. p. pp. when hind legs are bent at right angles to body. Eye very small. each with a 4-pointed "star" at the tip. 1925b. 104. Tins female has the skin on the back greatly distended by eggs. 2. 4. 1. or metacarpal tubercles present. 54. knee and elbow are widely separated. 56. heels slightly overlap. 6b. slightly webbed. 1. 1. fully webbed.— Rivero. chin without granules. regular tubercles surrounded by minute pustules. 2. a slight one at knee. Nostrils slit-like. which gradually descends to upper lip. their order from shortest to longest being 4. pp. 1937. 1958. 26. an adult female from Astillero. b2 2 88 Body very depressed and flattened. a heavy glandular lateral fold from behind axilla to groin. 18. venter minutely granular. 1959. c. Sabana de Mendoza. p. 277.— Mello-Leitao. no tarsal ridge.— Noble. its diameter about one-third its distance from tip of snout. 1948. 2. projecting. Tympanum not visible. Pipa parva Ruthven and Gaige. no skinfold across the chest.— Nic&oro Maria.— Barbour and Loveridge. pi. the second slightly longer than the third. 1927. — Miranda-Ribeiro. but with curving glandular ridges. fig. interval between nostrils about 2% times the snout length. p. with a few transverse glandular lines. figs. with pointed tips. no lateral skinfolds at angle of mouth. fig. 15. 14. p. no skinfold on heel. no true ventral disk. p. nearly equal in length. Snout long. p. size to 200 mm P.S. p. greatly depressed in profile and rounded at the tip. no inner or outer metatarsal tubercles. 38. 1961. Colombia. and larger flap-like appendages on tip of snout and at angle of jaw. p. 394. Canthus rostralis not distinguishable from loreal region.8 NATIONAL MUSEUM BULLETIN U.— A. upper eyelid not developed. Venezuela). — Carvalho. 8. Lutz. 8. Norte de Santander. Fingers long. 147122. a row of glandular ridges from posterior corner of eye to shoulder and along the sides. 1929. a few short tentacle-like skin structures near mouth and on throat. slightly projecting. palmar callus. the upper jaw extending far beyond the lower. Protopipa parva. p. 320. no thumb pad.

indicated only by a deep depression above angle of jaw. 1923b. upper eyelid not apparent. is is the one described above. Dunn The largest (1948) has given a very adequate discussion of relationships of this group. 9.— Gorham.5 mm. 8 —Head 11 AND GOIN 9 and body. west of El Mene. Tentacles on snout restricted to a few short pointed tubercles. 102 (type locality. as well as in Venezuela. no thumb pad present. tibia.. the upper jaw extending somewhat beyond the lower. as the whole head is flattened. 2. 1964. 19 mm. — 1914. Fingers free. . west of Rosario. the tips of the tubercles and the webs between the toes wood brown. 3.. hand. Dunn. Lagunillas. 1963. 507. edges of lips drab. 1925b. Eye very small. p. 15. 1. He notes the occurrence of this species in Rio Zulia. 1948. Specimens Examined COLOMBIA Norte de Santander: Astillero. Tympanum concealed. second the longest. p. mm.— Neiden. interorbital diameter three times the distance between the nostrils. Hno. Miiller. head width. 1914a. an adult from Leticia. 1 Pipa snethlageae L. Rfo San Juan. mm. pi. USNM 115819-20. femur. km. completely and those at angles minute. their tips with four small sharp projections. USNM 147122. MLS 270.FROGS OF COLOMBIA Dimensions. p. Brazil). 18. immaculate. fourth shortest and reaching to base of penultimate phalanx of third. 1924. p. 2. 4. — COCHRAN 18.— Noble. 1923.5 mm. of mouth not longer than eye diameter. and no palmar callus. very small. its diameter about one-fourth its distance from tip of snout. slit-like.. 293. Nic6foro Maria (1958) has figured and described some specimens from Astillero.. venter dull cinnamon to wood brown. north of Cucuta in the Department of Norte de Santander. Utinga. Toes pointed. p. two km. Snout very short. Colombia. p. bluntly rounded when viewed from above. and slopes out and then down to the upper lip. some drab mottlings between eye and shoidder below the glandular lines. Remarks. 275. pis. — Dorsal skin uniform olive. located on front and near center of snout. Para. figs.— Barbour. dorsolateral in position and on front of head. Amazonas. Pipa americana forma nova(?) Minke. first shorter than third. USNM USNM USNM 128845-6.5 mm. Canthus rostralis scarcely apparent.— Neill. individual —This known the smallest species of the genus.5 foot. —WTN Description 2561. 805-9. declivous in profile. Nostrils not projecting. near Belem. p. 45. Zulia. VENEZUELA: 15 115770-817. 115757-8. USNM 115767-9. p. 1914b. 419. 19. fig... Color in alcohol. head length. without lateral ridges. Pipa snethlageae Miiller Plate 1g-i 1914. p. metacarpal tubercles poorly developed. p. 9. 2. 2. mm.

a heavy glandular ridge above anus and two rounded projections below it. 89 mm. with small pale spots wherever the skin glands across chest. 1-5. venter very light gray with a dense scattering of darker gray and a few small black spots on center of throat and belly and along front of shoulder and forearm. p. except that larger.S. when hind legs are bent at right angles to body. 773. . 369. 92. 27. hind leg is adpressed. lightening to drab-gray on the sides and on legs. but no outer one. when limbs are laid along the sides. 1867. tubercles above eye and on sides of neck pearl gray. Skin of upper parts finely pustular. skin baggy on heel and knee without forming folds. no skinfold stout. Dumeril and Bibron. no tarsal Body very width of head. p. p. . Loose skin of body covers basal parts of legs and arms. in postaxillary region over 1% times the greatest When no external vocal sacs. Dimensions —Head and body. pi. 1948. second a little 2 88 shorter than fifth. with a slate-gray line from tip of snout below canthus to eye. the chin darker.5 mm. 1789. ending behind the depression above angle of mouth. a small blunt. indistinct glandular ridge from posterior corner of eye. — agrees well with Muller's description of the type. — occur. 3.— Dunn. 2. venter pustular. posterior femur slate-gray with lighter tips on the numerous granules and rows of tubercles there. p. a small round drab-gray spot immediately behind eye. with scattered glandular tubercles and short glandular ridges above eye and on neck and body. palms of hands and soles of feet clove brown. oval inner metatarsal tubercle.— Shaw.— Andersson. 9. femur. no ventral disk. the webs between the toes mottled with drab. life history. 1900. — Pipa americana Laurenti. becoming darker below eye.. 1. 13 mm. tibia.. 20. p. knee and elbow are widely separated. 210 (type p.10 U. Pipa pipa (Linnaeus) Plate 1d-f 1758. but no projection on tip of snout. p. pis. 1046. angle of jaw with a small pointed skin projection. — Boulenger. 167.. p. heel barely reaches axilla. head width. — Gimther. and its variation. head length.5 mm. ridge. p. Surinam). 35. more coarsely so on posterior abdomen and legs. 6. — Gmelin. 1854. 1768. More it is material of this interesting form should be sought in order to estimate occurrence. hand.5 mm.. the only specimen examined. and lightening and suffusing benind it. Color in alcohol Dorsum plumbeous. fig. — Steindachner. foot to end of third toe. Surinam). 51. no. side of head and body smoke gray. 20 mm. 31 mm. dots. Rana pipa Linnaeus. third toe longest. 1841. Remarks The example from Leticia. p. figs. 50. heels are separated. 1858.. 34. to end of third finger. 1802. a short. 25 (type locality. locality. NATIONAL MUSEUM BULLETIN webbed.— Wyman.

1948. 1 fig. p. 177. 1912. 1802.. — — 164. 1799. pi. 1923. — Fitzinger. 8.— Ruthven. Eye exshort. 112 of vol. 179 (type locality. Dunn. Pipa curucuru Spix. p. 1892. 65. making an obtuse angle when viewed from above. p. between the mouths of Rio Madeira and Rio Tapajoz). 1830. 1802. p.— Brandes 1830. p.— Crawford. 1841. p. dorsiger. 1 (type locality. 1826. 1. pi. flat-sided. 271. 104. 22.— Rivero. of plate facing 2. 1960. 1948. p. — Sonnini and (type localities. Dunn.— Goin. p. Latreille. p. p. p. 1 fig. Pipa pipa. p. Rio Pipa cururuu. 1948. Dunn. —MLS 810. 774. 78.— Barbour. 395.Ribeiro. p.— Baumann. 1948. 3. p. p. p. p.— Neill. figs. 434. p. 1901. p.— Rohl. 53. merging with the very narrow upper lip. Nostrils small.. Niceforo Maria. 86. Bufo p. Surinam). footnote 2 (type locality.— Ernst. Brandes and Schoenichen. p. 308. Negro. 2 (substitute for Rana pipa Linn. pointed in profile. 59. 1929.— Noble. pp. Barbour and Loveridge. 4. Lago Maximo. Description. Can thus rostralis not well marked. 1858. 13. 237. 1849. 156. p. 5-10. in Cuvier. p. p. 1960b. — — — 1948. pi. 90.— Miranda. 774. 1937. p. 1964. 1824. halfway between angle of jaw and tip of snout. 1802.— Boettger. Amazonas. 85.— Hno. and one-fifth of that distance in diameter. 214. p. p. p. 1S41. 1958. in Cuvier. p. 10. p. pis. 15. p. Pipa sedo Schlegel. Snout pi. 404. 1963. p. 96. 70. Durneril and Bibron. 4. Guiana and Surinam). lying at tip of snout. 14. 2.— Durneril and Bibron. 1835. p. 1963.— Sclater.— FROGS OF COLOMBIA — COCHRAN H AND GOIN 51— 1882a. Bahia(?) and 1826. fig. Pipa tedo Merrem.— Dunn. 199. Bartlett. Brazil. p. 39. 30. 5. 9. Ill. 1901. p. 2 (type localities. 1948. 1959. 1159. 1877. 1 fig. p. 9. p. 1923.— Tschudi. 1914. 1926. — Gravenhorst. p.— Rabb. 9. 34 (substitute for Pipa americana Durnenl and Bibron.). 1931. 121 1925. 95.— Dunn. Leptopus asterodactylus Mayer. p. Brazil) . 120. narrowly separated. 119. 1919.— Dunn. pi. — p. 31.— Rabb and Rabb. 10. 4. 1960. 22. Dumenl and Bibron. fig. 1895. 1836- Pipa 1836- 1948. loreal region narrow. p. 595. pipa. 774. p.— Wagler. Bancroft. 423.— Beebe. p. 155. p. laevis Duvernoy.— Gines. 417. p. 1. p. Colombia. Pipa pernigra Barbour. interorbital diameter nine or ten times that of the narrow upper eyelid. 105-110. Pipa 1820. 1948.— Cott. 1949. 1925. pi. 1923b. p.—Dunn. 10. 1962. 423.). p. tremely small. 172. figs. 122. figs. p. Rana dorsiger Schneider. 1923b. — — 1849. 320. 207. Pipa surinamensis Duvernoy. 194S. p. vertical. fig. Laur. 1961. 10. and Schoenichen. p. 1961. 1919.— Mello-Leitao. fig. fig. 40. p. Dunn.— Rabb and Snedigar.— Cochran. pi. 1896. p. 636. 101. 1858. p. figs. 1897. p. 275.— Dunn. fig. 2. p.— Nieden. 1833. p. 1870. an adult from Leticia. 1841. fig. 9. p. the upper jaw extending considerably beyond the lower. 1901. 1841.— Daudin. 17. near Villa Bella. p. 198. 281. 277. 1901. 774 — Brandes and Schoenichen. 2. p. p. p.— Minke. 10. photographs Bokermann. 1961.— Gorham. 1923. 17. 2. 1.— Gadow. 459. 1926. Aster odactylus fig. figs. p. 3 figs. 49. slightly projecting.— Schinz. Amazon River). pi. dorsigera.— Cope. p. and about three times that . 1838. p. 32. 28. 1829. 1960. — 1803. p.

Body very postaxillary region equal to greatest width of head. a small tentacle at each nostril. a female from British Guiana: head and body. BRAZIL: Rio Urubu. femur. but no outer one. near Itacoatiari. While Leticia. foot. with a transverse sepia line across the throat and a wider midventral stripe from throat to anus. triangular inner metatarsal tubercle. a sharp. 58. no pad. venter finely tubercular. 35 — USNM mm. but never produced into posteriorly directed epi- coracoid horns. 57571. — Specimens Examined COLOMBIA Amazonas: Leticia. is at present the only known Colombian locality for Pipa pipa. a narrow glandular ridge from eye to corner of mouth. or metacarpal tubercles. when limbs are laid along the sides. the second a little longer than thumb fourth. BRITISH GUIANA: MLS 810.. Dimensions. 54 mm. immaculate. not distinct.. tibia. no ventral no vocal sacs in the male. a slight skinfold stout and flat. disk. hand. Skin of upper parts roughened by small spiny tubercles beginning behind head and enlarging posteriorly. it seems likely that the species also will be found in headwaters of other affluents of the Amazon River system. SUBORDER DIPLASIOCOELA Shoulder girdle usually firmisternal. A young specimen (USNM 142178) from Rio Urubu near Itacoatiari. without lateral ridges. is ochraceous above and below. heel reaches axilla. Epicoracoids sometimes more or less separated. four longitudinal ridges on back and dorsolateral region. 57571. when hind legs are bent at right angles to body. with small sepia spots along the dorsal ridges and scattered over venter.. and a semicircular one above anus.5 mm.12 NATIONAL MUSEUM BULLETIN 288 U. each finger with a tactile 4-pointed "star" at its tip. 40 mm.. 134 mm.5 mm. in Amazonas. head length from angle of jaw. — Above dull olive. 53. Color in alcohol. palmar callus.. sides and venter with dirty whitish spots. third and fifth toes subequal. no skinfold across chest. 50 mm. knee and elbow are slightly separated. fully webbed. When hind leg is adpressed. and other tentacles nearly encircling eye. free.S. in on heel and knee. between Tympanum nostrils. Remarks. Fingers very long and slender. increasing in length in the order 1-4-2-3.. heels are well separated. USNM USNM 142178. Brazil. Toes long. head width. with many short transverse wrinkles on belly. Vertebral column either diplasiocoelous or uniformly . no tarsal ridge. a large fringed skinflap at angle of jaw.

D. 1 No a. Heel reaching to anterior border of eye. on underside of upper arm from axilla nearly to elbow. the sacral and presacral vertebrae are fused. size to 34. not over 18 . Outer metatarsals semi-ossified style. lugubris a. dorsolateral lines fine and sharp. Terminal phalanges T-shaped. no definite dorsal stripes. dorsolateral stripes relatively wide. a diagonal gland halfway between heel and inner metatarsal tubercle. size to 27. Family Ranidae Sacral diapophyses cylindrical or very slightly dilated. mm. pale axillary spot. D. D. Sacral vertebra usually articulated with urostyle by a double condyle. c1 Heel reaching a point between anterior corner . Generic diagnosis. . above. entire.FROGS OF COLOMBIA —COCHRAN AND GOIN 13 procoelous. D. No inter- calary cartilage between penultimate and last digital phalanx. . 1 . Tongue their tips dilated into disks subdivided Omosternum a weak. free or less distinct. size to mm 46 of eye. tinctorius Wagler). Skin usually smooth or faintly granular.5 mm. . sternum cartilagi- nous. elongate. size to 18 D. ternal.5 mm. Maxillary and vomerine teeth absent. 2 More or to Colombian Genera of Dendrobatinae Dendrobates maxillary teeth present less well-developed teeth on maxilla Phyllobates Genus Dendrobates Wagler Dendrobates Wagler. 1830. b2 of eye and nostril. Heel reaching posterior corner c2 . bl First finger longer than second. d : . Shoulder girdle firmis- pair of glandulomuscular organs present on top of each terminal digit. c1 A . Key a to Colombian Species of Dendrobates First finger nearly as long as second. Tympanum more behind. Subfamily Dendrobatinae Vertebral column uniformly procoelous. opisthomelas large white spot mm . Ribs never present. 2 First finger b1 much shorter than second. hahneli posterior half of belly only. A c1 . — Pupil horizontal. heel reaching to center of eye. trivittatus . 202 (type species. size to 23 No c2 . mm pale axillary spot. In Dendrobates. Key a. reticulations on D. p. . venter with light spots or reticulations evenly distributed. or longer. A Omosternum frequently bony. Fingers and toes free. united.. ingeri First finger nearly as long as second. Size small.

Dorsum pale. p. p. 7-12. fig. 1904. 858. p. usually a wide light middorsal stripe and a pair of dorsolateral stripes. projecting. 1963. . p. Dendrobates pictus (not of Dumeril and — Bibron). 1830. Description. pi. 8. Nostrils lateral.— De Witte. 1874b. Dorsum not black. 9. 2 (type locality. p. 1906. an adult female from Puerto Narifio. minutus minutus . 1933.— Beebe. 455. 273. p. 1 (type locality. snout very broadly rounded when viewed from above. Lutz and Kloss. 1901. . D.— Peters. p.— Parker. Tschudi.— Boulenger. p. — p. 92. irregular small black spots (in alcohol. Colombia. p. 18. b2. size to 37. 636. p. pi. p. 96. 202. 2. Ecgd. 71. 91. 1958. 1958. 181.— Lidth. 32. 1825. 448. Dendrobates obscurus Dumeril and Bibron. fig. 102. 1901. p. Lutz and Kloss. 94. 1939. Canthus rostralis . black with large or small light spots. 35. Teffe River.5 medium.— Crawford and Jones. 1825. 87. — Steindachner. tinctorius histrionicus size to 38 mm c2. p. 1952b. 87. 1961. 2a-d. 230.— Crawford. 13. p. figs.— 14 U. 9. 1864. 1885. p. p. 1884. Amazonas. p. c2 . p. 1942. 213 Cope.— Baumann. rounded in profile. photograph. size to 17. p. 665. p. D. — Dendrobates trivittatus . 1888d. 1824. 1945.— Gadow.S. Dorsum olive-gray with said to be scarlet in life) many . Cochran. NATIONAL MUSEUM BULLETIN Size to 38 mm. p. Brazil). 1931.— Miranda-Ribeiro. D. Guichenot. p. figs. up Dorsum c1 . their distance from end of snout one-half their distance from eye. ovate. 1830. fig. 98. p. Dendrobates nigerrimus. 669.— Mello Leitao. 669. large dark irregular spots. 1926. p. 1855. 257. fig. 1830. 2.5 mm. 1873a. size to 35. its posterior border free and unnotched. 1952.— Laurent.— Fitzinger. 148. p. minutus ventrimaculatus No conspicuous light spot below upper arm. p. — USNM 146832. 363. p. 129. D. tinctorius chocoensis d2 . D.— Andersson. 1841.— Schmidt and Inger. pp. Brandes and Schoenichen. Wagler. 164. Amazonas. 1913.5 mm.— Gorham. 112. size to 16. 655 (type locality unknown). 1882a. p.— Lutz and Kloss. the upper jaw extending well beyond the lower. 1843. tinctorius wittei Usually a large white spot on snout and another between shoulders. Hyla nigerrima Spix. 25. 1838.5 mm. p. 202. Hyla biviltata (lapsus for trivittata). 210. 1919. 36. p. 2. — 1952. 1912. pi. — Goin and Layne. . Many small white spots evenly scattered on back. .— Ahl. p. mm.— Kappler. 1937. 144. usually with d 1 . — — 1830. Brazil). p. 1885. pi. p. 161 Fowler.— Wagler. photograph. p. 1929. 1930. Boie (in Wied-Neuweid). d1 d2 . 404. d2 2 88 Heel reaching to posterior border of eye. Hyla trivittata Spix. 292. 1951. 37. tinctorius confluens Dendrobates trivittatus (Spix) Plate 5g-i 1824. — — 4. size to 32 mm. p. p. p. p. Tongue half as wide as mouth opening. D.

5 mm. separated from eye by an interval equal to about one-eighth its own diameter. Skin of upper parts granular. interorbital diameter 1% times that of upper eyelid. its diameter 1% times its distance from tip of snout.. 20. 17. their Color in alcohol. its tip reaching almost midway on penultimate phalanx of third. Tympanum distinct anteriorly.FROGS OF COLOMBIA — COCHRAN AND GOIN 15 slightly rounded. a pronounced skinfold on heel and knee. Eye large. venter with very indistinct granules and more distinct transverse glands across belly. 13.. throat and sides nearly smooth. disk of fourth toe covering less than one-fourth the tympanum. unspotted. knee and elbow overlap. side of head slate. posterior femur and post-anal region granular. and the dark slate color of its entire body when preserved contrasts with the Remarks. and continuing to groin. a skinfold across the chest..5 mm. — Dorsum blackish snout and anterior third narrow gray dorsolateral stripe running back from head. slate. — Dimensions. vertical. a swollen area behind tympanum partly obscuring its upper posterior border. heels overlap. head width. slender. venter and sides blackish slate. its greatest diameter about three-fifths that of eye. loreal region flat. first finger longer than second or fourth. —This one of the most readily recognized of the first finger is longer than the second a reversal of finger-length relationships found in other Colombian species except ingeri. a large palmar callus. equal to interval between nostrils. palms of hands and soles of feet olive-gray..5 mm. 17. foot. a tubercles and disks pale drab-gray. when hind legs are bent at right angles to body. femur. 43. a very small round thumb pad present. prominent. head length. with moderate disks and heavy lateral ridges.. third toe longer than fifth. Fingers rather long. 12 mm. Body elongate. Toes long. Head and body. when limbs are laid along the sides. a tarsal ridge ending in a diagonal bar and enlarged tubercle about two-thirds the distance from inner metatarsal tubercle to heel. Its back is granular. no glandular ridge behind eye. with a paler border along upper lip. a small oval inner metatarsal tubercle. and equal to that of distinct.. passing above tympanum. in postaxillary region narrower than greatest width of head. When hind leg is adpressed. the disk of which covers a little less than one-fourth the tympanum. free. its disk reaching halfway on antepenultimate phalanx of fourth. third finger.5 mm. heel reaches a point between eye and nostril. tibia. 11. in line with upper lip. and tarsometatarsal articulation reaches far beyond snout.5 mm.' frog is Colombian Dendrobates because the — . hand. metacarpal tubercles well developed.5 mm. less so on snout and just above dorsolateral stripes. no external vocal sacs in the male. of eyelids gray. and a smaller round outer one. a slight ventral disk.

continuous with the upper lip. Tympanum distinct but . 146847-9. 28931. size to 27. a small oval thumb pad present. pyriform.S. a small oval inner metatarsal tubercle. Body elongate. Para. Eye large. rounded. venter anteriorly slate-black. an adult from Aserrio. and a round outer one of about the same size. 24443. CAS PERU: Between Pucallpa and Rio Aquaytra. half as wide as mouth opening. Colombia. collected by Hno. USNM BRITISH GUIANA: Paruima MCZ USNM Dendrobates ingeri. a distinct skinfold on heel and knee. Can thus rostralis distinct. with a small round tubercle midway to heel. its posterior border free and unnotched. the upper jaw extending well beyond the lower. NATIONAL MUSEUM BULLETIN 2 88 pronounced patterns of the tinctorius group of Dendrobates.5 mm. Marajo Island. 127214-8. considerably narrower than interval between nostrils. 85152. the wide dorsolateral stripe appears as a lighter gray. Paratypes. Specimens Examined COLOMBIA Amazonas: Leticia. — rather small. posteriorly with faint light reticulations. truncate and sloping backwards in profile. same data as holotype. a larger round palmar callus. Putumato: Rio Aquiles on the Rio Mecaya. Orellana. Puerto Narino. USNM 144992-3. interorbital diameter equal to that of upper eyelid." MCZ USNM 15663. disk of fourth toe covering one-fourth the tympanum and appreciably larger than that of third finger. Toes long. ECUADOR: Junction of the Chiguancay and Chanchan Rivers. Caqueta. very slightly webbed at the base. Kamarang River. vertical. 146846. third toe much longer than fifth. their distance from end of snout one-third their distance from eye. however. 118681-2. phalanx of third. Niceforo Maria. near Rio Pescado. a weak tarsal ridge. projecting. its disk reaching to base of antepenultimate phalanx of fourth. with rather small disks and slight lateral ridges. Fingers long. metacarpal tubercles well developed. its greatest diameter one-third that of eye. BRAZIL: "Lower Amazonia. Mission. Nostrils lateral. its diameter \){ times its distance from tip of snout. snout bluntly rounded when viewed from above.16 U. Description of holotype. loreal region slightly concave. a tributary of the upper Rio Caqueta. Tingo Maria. Diagnosis. prominent. ANSP 18188. vicinity of Puerto Santa Clara. UF 8535. new species Plate 2d-f — USNM — USNM — Holotype. the disk of . First finger longer than second adpressed heel reaching to posterior corner of eye. free. separated from eye by an interval equal to one-third its own diameter. first finger longer than second or fourth. Sometimes. no dorsolateral light lines. Tongue relatively large. USNM 146832. its tip reaching to base of penultimate which covers about one-eight the tympanum.

slightly convex and sloping backwards in profile. projecting. Skin of upper parts coarsely granular except on snout. continued on upper proximal surface of upper arm. 11 mm. Dendrobates hahneli Boulenger.. when limbs are laid along the sides. trivittatus in the key. its diameter about 1% times its distance from tip . and a wide glandular ridge from posterior corner of eye descending behind tympanum and ending above shoulder. 10 mm. with a few fine. 613. their distance from end of snout about one-quarter their distance from eye. reticulation posterior — Remarks. Canthus rostralis distinct. and with traces of a gray chevron mark in front of eyes. knee and elbow touch. venter smooth (pustular under the lens). — Dorsum slate-black. head width. Colombia. up to 27. palms of hands and soles of feet slate color. no skinfold across the chest. 8 mm.FROGS OF COLOMBIA — COCHRAN AND GOIN 17 in postaxillary region a little narrower than greatest width of head. Dendrobates p(ictus) hahneli. 412. northern Peru) 1886.. and its adult size is larger. 12 mm. 8. Head and body.5 mm. a small round light spot in axilla. a small pearl gray spot on upper proximal femur behind groin.5 mm. Description. Although the possession of the long first finger places D. 601. prominent. 1952. apparently no external vocal sacs in the male.. Dendrobates hahneli Boulenger Plate 5d-f 1884. p. venter slate-black. Dimensions. while its first finger is clearly longer than the second. no ventral disk. and a light stripe around edge of lower jaw. Meta. heel reaches posterior corner of eye. 636. heels slightly overlap. the area along the upper jaw mouse gray. 57. with very indistinct indications of a coarse light on belly. tibia. Color in alcohol. head length. 3. an adult from the the upper Giiejar and El Mico. 4. femur slate-black. unspotted. The new species has a much shorter hind leg and grows only half as hahneli. a little lighter on top of head. 4 (type locality. a slight swelling behind eye. B. pi. 1884a. their tubercles and disks drab-gray. when hind legs are bent at right angles to body. irregular glandular lines near the sides. the upper jaw extending well beyond the lower. ingeri with D. fig. merging with the vertical upper lip. on Tongue half as wide as mouth opening. its posterior border free and unnotched. — — USNM 144965. the two are not closely related. figs. 1952. Nostrils inferolateral. This new species apparently is allied to Dendrobates but it lacks the dorso-lateral light lines.. which is nearly smooth. side of head slate-black. vertical. Yurimaguas.. Huallaga River. loreal region flat. p. foot. pp. hind leg is adpressed. . ovate.5 mm. Macarena. Lutz. femur. large. When — 7. Eye large. hand. 27..5 millimeters. snout truncate when viewed from above.

18 U. When hind leg is adpressed. 21 mm. the pale axillary spot does not exist . crossing lip diagonally and ending above shoulder. side of head black. foot. —The structure and markings of this form are rather like those of Dendrobates lugubris. with lateral ridges. Color in alcohol. Skin of upper parts finely granular. the disk of indicated by lateral skinfolds at sides of chin. with a fine gray reticulation on and a gray patch on lower inner halves of femora united on the midline. Toes long. when limbs are laid along the sides. with some coarse granules on sacrum. its tip third. a glandular swelling from posterior corner of eye extending across upper posterior part of tympanum and ending behind it. a similar pearl gray spot in axilla. The adult D. venter smooth. a small weak thumb pad present. a distinct oval inner metatarsal tubercle and a smaller round outer one. a distinct skinfold on heel and knee. soles of feet and palms of hands black. — Dimensions. with a tubercular swelling midway.. and lightening and below the knee. its greatest diameter half that of eye.. tibia. a blunt tarsal ridge.5 mm. no ventral disk. interorbital diameter 1% times that of upper eyelid. 9 mm. 6. head length. where it ends in a large oval pearl gray spot on upper anterior femur. hand. knee and elbow touch. head width. fourth longer reaching to base of penultimate phalanx of which covers about one-eighth the tympanum. and another one on concealed surface of tibia behind eral line beginning at eyelid snout tip. but numerous differences can be noted. first finger nearly equal to second. with a few fine diagonal striations on proximal part of posterior femur. disk of fourth toe covering one-eighth the tympanum and approximately equal to that of third finger.5 mm. 6 mm. a more definite round palmar callus. with a narrow slate-gray dorsolat- continuing on canthus and upper to pearl gray on the side. femur.. 8.. —Dorsum black. no skinfold across the chest. hahneli is smaller. with a fine pearl gray line beginning as a small crescentic mark on tip of snout. ending at heel. heels slightly overlap. Remarks. their disks and tubercles posterior half of belly lightening to drab-gray. third toe longer than fifth. when hind legs are bent at right angles to body. not separated from eye by an appreciable distance. Body elongate. and above tympanum. and behind anus. heel reaches center of eye. Fingers long. venter black. 10. Tympanum distinct except for its upper posterior edge. Head and body. external vocal sacs in the male than either. its disk reaching to base of antepenultimate phalanx of fourth. touching lower edge of nostril.. 7 mm. greater than interval between nostrils. on upper surface of legs. metacarpal tubercles well developed.. free. in postaxillary region a little less than greatest width of head.5 mm. continuing on lower surface of upper arm to elbow. NATIONAL MUSEUM BULLETIN 2 88 of snout.S.

1963. p. 1952. 1917. in northern Peru. slightly projecting. fig. vertical. Boulenger. and shore of Guayabero. fig. lugubris. slightly more than one-quarter as wide as mouth opening. 5. Tongue very slender. Apparently this species is the only one of the genus found east of the Andes in Colombia. and in hahneli the light ventral reticulations are confined to the posterior half of the lines are much finer and sharper than abdomen and the dorsolateral in lugubris. p. — — Stebbins and Hendrickson. More material from both countries is needed to determine whether truly significant differences in bodily proportions might warrant separation. 144950-8. the upper jaw extending well beyond the lower. 10.— Dunn. its posterior border free and unnotched. an adult from Mariquita. Boulenger stated that the snout and eye were equal in length and that the interorbital width was greater than that of the upper eyelid. ham. Domo Meta: USNM USNM CNHM USNM Santa Clara. Phyllobates lugubris. (CNHM The Putumayo frog 61789) has a snout length equal to 1% times the eye diameter. 127201. —USNM Description. — Dunn. pi. 1882a. 1959. 502. 250. truncate and sloping backwards in profile. p. 134. snout nearly truncate at the tip when viewed from above. Dendrobates lugubris O. Dendrobates lugubris 0. pp. USNM 127932. In his original description. near Camp 1. p. 144962-7. ovate. 1944c. it is not feasible to consider a separation on these extremely variable characters. while the interorbital width is broader than that of the upper eyelid. While the first finger usually is slightly shorter than the second in both are equal in length in about 10 percent of the examples hahneli. Until more Putumayo specimens are examined. 61789. Macarena. PERU: Orellana: Campo Santa Clara.— FROGS OF COLOMBIA in — COCHRAN AND GOIN 19 D. 2. Colombia. 145. 14 (type localities. 132. their distance from end of snout about half their distance from eye. — Gor- 25. 144959-61. and the examples from Meta and Putumayo seem to agree well with the description of Boulenger's type from Yurimaguas. 1924. in the Macarena. p. 146850. 1924. at hand. Putumayo: Umbria. 127203-8. New Granada and Costa Rica. Schmidt Plate 2a-c 1858. Specimens Examined COLOMBIA USNM Cafio No. p. p. descending to the vertical . Guayabero River shore above El Refugio. Tolima. Baumann. Nostrils lateral.— Taylor. loreal region flat. Canthus rostralis rounded. 520. Upper Giiejar and El Mico. on road between Boca del Toro and Chiriqui Volcano). 2. 639. Schmidt.

10 mm.. venter nearly smooth. 10 mm. with a slate-gray dorsolateral stripe covering the gland from tip of snout to groin. a heavy skinfold on heel and knee. Dorsum black. and forearm similarly spotted. a very one-third the small round thumb pad present. the tubercles juveniles (USNM — As there are no traces of maxillary teeth in the numerous Colombian . its diameter equal to its distance from tip of snout.. its greatest diameter three-quarters that of eye. with faint lateral ridges. lower limb surfaces similarly marked. foot. a ventral disk.5 mm. Eye large. This species has been considered in the genus Phyllobates by Dunn (1924) and by Taylor(1952). posterior femur with large irregular gray spots on a black ground. prominent. on posterior abdomen. Skin of upper parts finely granular. which equals fourth. a faint tarsal ridge. with a gray crescentic mark below eye. Remarks. ending on knee. distinct. the chin sometimes citron yellow. 13 mm. interorbital diameter twice that of upper eyelid.20 U.. hand. a few fine skinfolds across the chest and throat. its anterior border practically in contact with eye socket. and sides chromium to apple green. Color in alcohol. palms — — and disks drab-gray. above eyelid and along sides to groin. head width. no supratympanic ridge. the tympanum and nearly the same size as that of third finger. Toes moderate. femur. disk of fourth toe covering one-third large palmar callus. its disk reaching halfway on antepenultimate phalanx of fourth. 147159-63) have light areas on back. 12 mm. Dimensions. although Dunn (1944c) later reversed his opinion by restoring it to the genus Dendrobates. light areas of belly olive-yellow. knee and elbow overlap. with a few pustules on chest. heels overlap. free.. when hind legs are bent at right angles to body. of hands and Some soles of feet black. Head and body. a little greater than interval between nostrils. 31. the disk of which covers about tympanum. a small oval inner metatarsal tubercle and a round outer one of about the same size. tibia. and below anus. sides of head black. 14. in postaxillary region narrower than greatest width of head. tibia. when limbs midway are laid along the sides.S. 10 mm.. a metacarpal tubercles well developed. ending in a short diagonal gland to heel. tips of second and fourth fingers reaching to base of penultimate phalanx of third. and an elongate irregular gray spot on outer half of femur. heel reaches center of eye. a large roundish gray spot on proximal part of anterior femur. first finger a little shorter than second. third toe longer than fifth. Fingers long. When hind leg is adpressed. Tympanum large.5 mm. apparently no external vocal sacs in the male.. foot upper arm. a wide flat dorsolateral gland from tip of snout. NATIONAL MUSEUM BULLETIN 2 88 upper lip. venter black with wide gray crossbands or spots. Body rather elongate. head length.

however. 100 m. 61181. 122104. Magdalena: Santander: USNM Rio Frio. west of Playa de Oro. CNHM Espinal.. USNM 150168. north of Alban. Uraba." USNM 48865.. rarely. Dunn previously had identified 48865 from Truando as "auratust. Cundinamarca. the light markings no longer are apparent. its — COCHRAN AND GOIN position in the genus Dendrobates 21 is accepted here. PANAMA: Almirante. 150687-91. Caparrapi. Mariquita. 54200. other examples still has traces of primrose yellow and on the light dorsolateral lines (USNM light areas on sides and 145099-100. USNM USNM 144968-9. USNM 144944. the respective measurements of the smallest one are 6.5 millimeters. Caparrapi. in preservative. Three tadpoles (USNM 144985) that possibly are of this species were collected at Volcanes. The adpressed heel may reach to the center or the posterior corner of the eye or. Finca La Esmeralda. ANSP 25763. near Pep6." ANSP 2722-3. Mesa Cucuana." but that specimen appears to be more like the all- USNM black lugubris. Rio Currulao. Dunn on October 20. "Truando. Villa Arteaga. MLS 387. Simacota. shore of Rfo Guali. Santa Cecilia. ANSP 21030. 145102-3) are pale Nile blue in those areas. and extreme limits of llanos. The first finger is slightly to distinctly shorter than the second in nearly all of the adult examples at hand. Choc6: Bahia Utria. Tolima: 144974-7.FROGS OF COLOMBIA specimens at hand. 530-550 m. ANSP 24324-31. USNM CNHM 63878. "New Granada. Santuario. La Victoria. by E. Ungufa. USNM 103. Such examples closely resemble D. near Lebrija. Cundinamarca: East base of Cordillera. upper Rio San Jorge. near Lebrija. 1943. the long first finger in trivittatus and the much shorter one in lugubris should serve to differentiate the two.5 and 9. 81870. Specimens Examined COLOMBIA Antioquia: CNHM 54275. One specimen on the (USNM 145098) jaws. 10-15 mi. Finca 118223. Golfo de Uraba. which also is often very dark after preservation. MCZ 16059-64.. and the whole body appears slate-colored or blackish. 63877. R. Mariquita. Caldas: Pereira. USNM 144971-3. USNM USNM 145095- 144978-9. Sometimes. USNM 147159-63. Rio San Juan. The largest one measures 7 centimeters in head and body length and 11 centimeters in tail length. 144970. 120 m. Chicoral. . 144948. Pueblo Rico. Cauca River. 337-262—70 3 USNM 118231. trivittatus. El Mosco. probably of this species). 81826-8. only to the tympanum. 146850-6. Volcanes. USNM 98891-3. El Centro. MLS 314-5. CNHM Catival. USNM 144985 BoLfvAR: CNHM CNHM CNHM (three tadpoles. USNM Nechi.

prominent. foot. loreal their region flat. When hind leg is adpressed. no true tarsal ridge. rostralis than interval between nostrils.2. Tongue relatively large.. in line with the vertical upper lip. Fingers moderate. its tip reaching to base of penultimate phalanx of third. belly finely granular. north of Medellin.30. a swelling behind eye.800 ft. BM USNM 145794 (formerly Description.5 mm. hand. lateral. Eye large. 18 mm. fig. no skinfold across the chest. Dimensions. apparently no external vocal sacs in the male. coan adult from Santa Ines.. vertical. north of Medellin.. Skin granular on back and sides. — Gorham. pi. ovate. Head and body. its diameter a little greater than its distance from tip of snout. and a distinct glandular ridge beginning behind eye and descending behind tympanum. tibia. head length. when hind legs are bent at right angles to body. when limbs are laid along the sides. in postaxillary region a little wider than greatest width of head.5 mm. 7 mm. p.third broadly rounded. p.. Santa Inez. ending on shoulder. Colombia.15. rounded and sloping backwards in profile.. Nostrils type).S. 4 (type locality. 275. disk of fourth toe covering half the tympanum and approximately equal to that of third finger.. 25. Antioquia. NATIONAL MUSEUM BULLETIN 2 88 Dendrobates opisthomelas Boulenger Plate 2g-i 1899.22 U.— 1947. a strong skinfold on heel and knee. 3. a larger metacarpal tubercles well developed. covering its upper posterior rim. 8 mm. the upper jaw extending well beyond the lower. Antioquia. Body moderate. 5. Andes of Colombia).5 — mm. but a diagonal gland midway between heel and inner metatarsal tubercle. 5. 11. third toe much longer than fifth. Tympanum distinct except on its upper posterior border. a faint ventral disk. first finger much shorter than second. slightly eye. the disk of which less covers half the tympanum. Toes long. femur. snout broadly rounded when viewed from above. knee and elbow greatly overlap. fourth a trifle longer than second. with faint lateral ridges. chin and chest smooth. Dendrobates opislhomeleas Boulenger. callus. their distance distance from Canthus from end of snout one. heels slightly overlap. especially so on sacrum. its posterior border free and unnotched. 6 mm. . head width. rounded palmar a small oval thumb pad present. separated from eye by an interval equal to one-third its own diameter. a small inner metatarsal tubercle and a smaller but equally distinct outer one. 1963. free. its disk reaching just beyond base of antepenultimate phalanx of fourth. its greatest diameter about one-third that of eye.. interorbital diameter 1% times that of upper eyelid. slightly projecting. head nearly smooth. heel reaches anterior corner of eye. three-fifths as wide as mouth opening. 7.

palms of hands and soles of feet drab. snout nearly truncate when viewed from above. —Dorsum — COCHRAN seal AND GOIN 23 brown. and some smaller and less distinct light spots on anterior proximal femur and in groin. side of head drab. these spots extending on the sides where they darken to merge with the dorsal color. the disk of which covers about three-fourths the tympanum. near Rio Pescado. and belly. 147224. the upper jaw extending well beyond the lower. scarcely separated from eye. Dendrobates minutus ventrimaculatus Shreve Plate 3d-p 1935. Fingers moderate. Toes moderate. Eye large. 1958. third toe slightly . free. loreal region flat. 5194-6. Tongue a third as wide as mouth opening. p. although hahneli and minutus ventrimaculatus approach its lowest proportions. Tympanum distinct. vertical. their distance from end of snout about a fifth their distance from eye.15. p. interorbital diameter 1% times that of upper eyelid. This minute frog seems to be confined to Antioquia. a large pale ecru drab spot on underside of upper arm from axilla nearly to elbow.800 ft. continuous with upper lip. their disks and tubercules a little lighter. The heel reaches to the anterior corner of the eye in opisthomelas. a small round palmar callus. Nostrils lateral.30] (cotype). but not beyond its posterior corner in minutus ventrimaculatus. metacarpal tubercles well developed. CNHM USNM 63874-5. with lateral ridges. Rfo Negro. Santa Ines. AMNH [formerly Llano Grande. 147506-7.2. a little less than interval between nostrils.— Marx. — Remarks. 213 (type locality. 1947b. north of Medellfn. prominent. USNM USNM BM 1947. Caqueta. Dendrobates minutus ventrimaculatus Shreve. with a pale stripe along upper lip border from tip of snout extending to shoulder. 431. chest. Specimens Examined COLOMBIA Antioquia: Medellin.FROGS OF COLOMBIA Color in alcohol. Canthus rostralis rounded but distinct. 145794 AMNH 1372-4. slightly projecting. venter sepia. Colombia. an adult female from Aserrio. 3. a small oval thumb pad present. — Description. truncate and sloping backwards in profile. its greatest diameter half that of eye. 1364. first finger much shorter than second. Sarayacu» Ecuador).. The head appears to be both longer and wider in relation to total length than in any other Colombian species of Dendrobates. upper limb surfaces sepia. fourth longer than second and reaching to base of penultimate phalanx of third. its posterior border free and unnotched. with large light ecru drab spots on chin. its diameter equal to its distance from tip of snout. oval. probably Medellm. USNM 146843. p. 316.

and smaller than that of third finger. when hind legs are bent at right angles to body. a slight ventral disk. . with a rather narrow drab-gray Color in alcohol. Dorsum black. MLS 42a is very similar to the described specimen but has several black spots across the throat instead of a single wide black band. the tips of the digits lightening to drab gray. a weak skinfold across the chest. with a few indistinct gray transverse spots on tibia. when limbs are laid along the sides. with several large round black spots on belJy. in postaxillary region equal to greatest width of head. . The frog from the watershed of Rio Pastaza. Skin of upper parts smooth (pustular under the lens) venter minutely granular. ridge. 16 mm. . sides of body black. with a gray line along upper lip border. 5 mm. a wide black band across throat. femur black near the body. Remarks.. disk of fourth toe covering about a third the tympanum. and ending on side. a short median drab-gray line beginning between so. 6. a — . a distinct tarsal with a small crescentic swelling near its center. a pair of large lateral black spots on chin. minutus ventrimaculatus. tibia. very fine glandular ridge from posterior corner of eye. Ecuador. In all six.5 mm.. dorsolateral line starting on snout. The heel reaches to the center of the eye in three individuals. the throat and chest less . 4. soles of feet and palms of hands black. heel reaches to posterior corner of eye. with an indistinct gray lateral strip. continuing along canthus and eyelid on the side to groin and ending on anterior femur in a wider irregular gray spot. Although we do not have enough specimens to make an adequate variational study of this subspecies. knee and elbow touch. Body moderately elongate. NATIONAL MUSEUM BULLETIN 288 fifth. sides of head black. Dimensions. femur.S. above tympanum. its disk reaching to center of antepenultimate phalanx of fourth.24 U. head length. becoming paler on distal half.5 mm. When hind leg is adpressed. heels touch. venter drab-gray. behind the knee and on proximal one-third of tibia. a skinfold on heel and knee. and to its anterior border in one example. 5 mm. to its posterior border in two specimens. a distinct oval inner longer than metatarsal tubercle and a smaller round outer one. the first finger is much shorter than the second. reported by Andersson (1945) as Dendrobates tinctorius Schneider undoubtedly is D. 7 mm. head width. 7 mm. limbs black above. Head and body.. a few facts about the individuals at hand may be mentioned. as Andersson's color desposterior this pale area continuing — cription agrees with the original description of this subspecies as well as with the specimens at hand. foot and forearm. apparently no external vocal sacs in the male. hand. forking anteriorly to join the pale canthal line in front of the eyes and ending between the shoulders. — the eyes. and a small median black spot on tip of chin. ending at heel. foot.

its tip reaching to base of penultimate phalanx of third. Antioquia. a round palmar callus. collected by Hno..thirds the tympanum and smaller than that of third finger. Can thus rostralis bluntly rounded. 54223. USNM Aserrio. no true tarsal ridge. vertical. Nostrils lateral. Dendrobates minutus minutus Shreve. 25. Barro Colorado 1963. Dendrobates minutus minutus Shreve Plate 3a-c 1958. Nic&oro Maria) from Sutatenza. Tympanum distinct. Fingers moderate in length. Eye large. Body moderately elongate. are quite similar to those mentioned above but differ from them in being much lighter in color. their distance from end of snout being half their distance from eye. near Rio Pescado. p. its diameter about 1% times its distance from tip of snout. but a short diagonal gland crossing tarsus midway to heel. 212 (type locality. its disk reaching to center of antepenultimate phalanx of fourth. No maxillary teeth can be found. a distinct skinfold on heel and knee. snout broadly rounded when viewed from above. fourth a little longer than second. 54203. truncate and slanting backwards in profile. Oriente: Rio Pastaza. equal to interval between nostrils. third toe longer than fifth. USNM 118705. Tongue almost La Ceja. Toes moderate. heel reaches to posterior corner . continuous with the upper lip. 1963. prominent. an indistinct thumb pad present. an adult from Diablo. Panama). with traces of a web between the three outer ones. loreal region flat. Antioquia. slightly projecting. Colombia. free. metacarpal tubercles well developed. When hind leg is adpressed. in postaxillary region a little narrower than greatest width of head. oval. first finger much shorter than second. with faint lateral ridges. the upper jaw extending slightly beyond the lower.FROGS OF COLOMBIA — COCHRAN AND GOIN 25 Five examples (USNM 146834-8. Description. disk of fourth toe covering two. they might be grouped with one of the smaller kinds of Phyllobates. Specimens Examined COLOMBIA Caldas: Caqueta: ECUADOR: Santa CNHM Pueblo Rico. Monte del mouth half as wide as opening. interorbital diameter a little greater that that of upper eyelid. Island. p. — USNM — Gorham. its greatest diameter about one-third that of eye. otherwise. 146843-5. separated from eye by an interval equal to one-quarter its own diameter. Dendrobates minutus. m. its posterior border free and unnotched. all the finger disks relatively large. an indistinct inner metatarsal tubercle and a poorly defined outer one. the disk of which covers about three-quarters the tympanum. 500 Cecilia. 144980.

— — Dendrobates tinctorius (not of Schneider).. when hind legs are bent at right angles to body. the chest. head width. 1923a. Breder. 6 mm. tibia. their disks and tubercles a little lighter. Los Mangos.5 mm. with a wide drab band across throat and a small drab spot near tip of chin. knees touch. 7 mm.26 U... 6 mm. an adult from one kilometer west of . and elbow. ANSP 23126.. USNM 144980-4. 17 (type locality. photograph. femur. 17. Dendrobates tinctorius wittei Laurent. unspotted. Of four tadpoles (USNM 144984) that were collected with the La Ceja frogs. 5. knee and elbow touch. minutus from Barro Colorado Island. they agree with the topotypes of minutus in all other essentials. fig. completely free.S. hand. Color in alcohol. — USNM 124223. The appearance of minute hind legs on the latter tadpole shows the beginning of metamorphosis. — Dimensions. the actual range in color pattern may be found to be quite similar. While the La Ceja frogs are darker and show less pattern than typical D. soles of feet and palms of hands sepia. Remarks. p.5 mm. and NATIONAL MUSEUM BULLETIN 288 tibio tarsal articulation reaches to tip of snout. lightening to sepia near edge of upper jaw. except top of head. as well as the fingers. Description. head length. in which most forms have the rior — toes. the smallest has a head and body length of 6 millimeters and a tail length of 13 millimeters and the largest has respective measurements of 9 and 16 millimeters. Barbour. with a small lighter sepia spot on knee. 1927a. Gatun. postefemur clove brown. MCZ 15288 (holotype). When more examples from both localities have been collected and studied. ANSP 21441-4. which is smooth.5 mm. Panama. —Dorsum clove brown. a slight ventral disk. when hind limbs are laid along the sides. 72.. 7. Barro Colorado Island. The faint traces of webs between the outer toes in this subspecies is unusual in the genus Dendrobates. 23200. of eye. venter seal brown. 4. p. PANAMA: Dendrobates tinctorius wittei Laurent Plate 5a-c 1923. heel. venter smooth (pustular under the lens) a pronounced swelling behind eye and a wide glandular ridge from posterior corner of eye descending behind tympanum and ending above shoulder. 1942. Specimens Examined COLOMBIA Antioquia: La Ceja. 13. sides of head clove brown. no skinfold across . 12. Skin of upper parts faintly granular. p. no external vocal sacs apparent in the male. [Magdalena?] Colombia). Head and body..5 mm. Monte del Diablo. foot.

first finger shorter than second. with small disks. in postaxillary region smaller than greatest width of head. smaller on anterior parts.. Tongue small.5 mm. femur. snout wide at the tip and nearly truncate when viewed from above. free. . vertical. no apparent thumb pad or palmar callus. 12..5 mm. Choco. edges of lower lips with small black dots or spots. but instead a glandular swelling suggesting a parotoid gland. some black areas on lower limb surfaces corresponding to position of dorsal silvery crossbands. venter pearl-gray. foot. Skin of upper parts smooth (under the lens appearing minutely pustular everywhere) venter minutely pustular. fourth equal to second. slightly projecting. Eye large. Colombia.5 mm.. no glandular ridge behind eye. disk of fourth toe covering about one-fourth the tympanum and much smaller than that of third finger. metacarpal tubercles scarcely evident. tibia. vocal sacs. Toes moderate. with enlarged disks. the upper jaw extending well beyond the lower. with an irregular black patch on center of throat. black dots. —Black many rounded silvery-white much larger on sacrum. head width. no tarsal ridge. 15 mm. interorbital diameter 1% times that of upper eyelid and a little greater than interval between nostrils. upper lip edged with . Fingers moderate. knee and elbow overlap when hind legs are bent at right angles to body. with a few small flattened granules on posterior part of belly and below vent. Nostrils inferolateral. ovate. Color in alcohol. its greatest diameter half that of eye. with one large white spot and two smaller ones taking the position of the usual diagonal crossbars. heel reaches to posterior corner of eye. separated from eye by an interval equal to about one-third its own diameter.. no skinfold across chest. no ventral disk. a weak oval inner metatarsal tubercle and a very small outer one. Tympanum fairly distinct... When hind leg is adpressed. 10. third toe longer than fifth. their distance from end of snout a little less than their distance from eye. prominent. hand. loreal region slightly concave. its disk reaching halfway along antepenultimate phalanx of fourth. their tips reaching just beyond base of penultimate phalanx of third. 35. 14 mm. with a large white spot on both upper and lower arm. descending straight to the upper lip. heels barely touch. no external its .FROGS OF COLOMBIA —COCHRAN AND GOIN 27 Andagoya. upper surfaces above. one-fourth as wide as mouth opening. without lateral ridges. with spots. its diameter equal to distance from tip of snout. a prominent skinfold on heel and knee. Body moderate.5 mm. — Dimensions. region around anus black. 10 mm. when limbs are laid along the sides. head length. Canthus rostralis bluntly rounded. arms black above. 10. the disk of which covers nearly half the tympanum. flat and slanting backwards in profile. sides white with a few of legs black. Head end body. its posterior border free and unnotched.

their distance from end of snout about two-fifths their distance from eye. as traces of that color still remain on some other individuals of this lot. 2-4. t.— Gorham. Andagoya. t. figs. NATIONAL MUSEUM BULLETIN 2 88 black. 144991. fig. (type localities. 13707-9. 12.S. Rio San Pablo.90-3. Quito. running south and west to the Pacific. Finca La Victoria. 147510. p. D. p. Granada). the upper jaw extending well beyond the lower. p. 13703-5. 76. loreal region flat. 1913. 1944c. 520. 3 figs. Choc6. Upper Condoto. Boulenger. USNM Andagoya. is found from the seacoast west of the Western Cordilleras. 1963. 104. p. 147508-9. wittei.. 13716-9. 1.12. D. species occurring in the — the Atrato. and the Baud6. Another. its posterior border free and unnotched. along the Rio Pepe to Andagoya and somewhat to the east. 1915. the silvery. New 1845. 43. 1944.] Dendrobates tinctorius histrionicus Berthold Plate 4g-i 8 (type locality. oval. 1027. Description. vertical. 1956. more directly to the Pacific. Dunn.58-61.21. Specimens from Sierra de Baud6 have been classified as probable intergrades between for three rivers D. p.11. chocoensis. 1913. tinctorius histrionicus. Bahfa de Utria. near Pepe" on shore of Rio Baud6. p.— Funkhouser. appears to be only from the vicinity of Andagoya. Specimens Examined COLOMBIA One km. One form. upper Rio 144986-90. BM USNM 145791. an adult from 75 miles south of Andagoya. the San Juan. 13695-701. t. — — — USNM figs. Tongue almost half as wide as mouth opening. pi. D. 1915. 1864. Can thus rostralis distinct. 1942. Nostrils lateral. Dendrobates tinctorius cocteaui.10. Boca de la Raspadura. 12. Dendrobates histrionica. p. in the valley between the Central and the Western Cordilleras. Laurent.000 ft. . a large white spot on loreal region extending below eye and another on tip of snout. 261 pi. The third one. slightly projecting. Dendrobates tinctorius histrionicus. Dendrobates tinctorius cocteani (lapsus. 1942.white spots were rich geranium-pink. (In life. comes from inland and southward.21. flat and sloping backwards in profile. wittei and D. snout truncate when viewed from above. [Probable intergrades: wittei USNM USNM BM AMNH X USNM ANSP 25617-25. Colombia. t. but intensive collecting decide the advisability of putting wittei into the is needed to synonymy of the latter. chocoensis. AMNH 13711-2.28 U. Brazil and Colombia).) —The anomaly of possibly three subspecies of a single Choc6 can be explained by plotting their distribution on a map. flowing west. for cocteaui) Steindachner.63-5. west Choc6: of USNM 124218-32. running north to empty into the Caribbean. Near Andagoya are watersheds Remarks. — 139778. 25. 1913. Sierra de Baud6. Dendrobates histrionicus Berthold.10. chocoensis. upper bank of Rio San Pablo.

tips of its second and fourth fingers reaching to base of penultimate phalanx of third. 12 mm. palms of hands and soles of feet brown. first its by an greatest diameter interval equal to own diameter. 13. foot. finger shorter than second. head length. no tarsal ridge. head width. often covering the whole snout anterior to them. Toes moderate. however. This form. 29 its diameter distance from tip of snout. with an oval white spot on posterior surfaces of femur and tibia and on top of foot and forearm. with a large round white spot encircled by a darker border on center of back. venter dark sepia. Skin of upper parts smooth (minutely pustular under the lens) venter smooth. prominent inner metatarsal tubercle. femur. Fingers moderate. Head and body. Body elongate. no ventral disk. a very weak thumb pad present. upper lip white from eye forward. as there is usually a large light spot between the eyes. and another on the sacrum.. 10. Sometimes a large round or oval light spot occurs on the back above the shoulders. the disk of which reaches almost to base of penultimate phalanx of fourth. Eye large. when hind legs are bent at right angles to body. without lateral ridges. interorbital diameter a trifle greater than that of upper eyelid.5 mm. no glandular ridge from posterior corner of eye. a small. tibia. Often. FROGS OF COLOMBIA in line with the vertical equal to its upper — COCHRAN lip. knee and elbow touch. 10 mm. whose Latin name means "relating to acting. no skinfold across chest. AND GOIN prominent. Dimensions. when limbs are laid along the sides. or the entire back may be light.. When hind leg is adpressed. a specimen in which the venter is light except for a black — — — USNM ." is another of the frogs having a clown-suit pattern. in postaxillary region narrower than greatest width of head.. limbs wood brown above. metacarpal tubercles scarcely indicated. third toe longer than fifth. 10 mm. two-third that of eye. the tip of which covers half the tympanum. with a large light spot on chin and chest and another one covering the posterior half of the abdomen. no external vocal sac apparent in male. separated from eye one-third free. the disks a little paler. Dorsum wood brown. rest of head brown. but a slight swelling in parotoid area. and a smaller round outer one. equal to interval between nostrils.. round.5 mm.. heel reaches posterior corner of eye. 33. Tympanum distinct except on posterior rim.. An exception is 144945. a skinfold on heel and knee. which is equal to fourth. Remarks. heels touch. hand. including lower anterior snout. with a large rectangular dark-bordered white spot covering throat and chest and another on the posterior part of belly. 14. This pattern is somewhat more regular than in Dendrobates tinctorius chocoensis. disk of fourth toe covering about one-fourth the tympanum and much smaller than that of third finger. a weak palmar callus. Color in alcohol.. the belly is black.5 mm.5 mm.

S. as the frogs did not die simultaneously. 30 km. shoulders. These differences may be due to the progressive state of decom- position before preserving. In the most-spotted individual the back was gallstone yellow where not black. 188. t. chocoensis in color pattern. and legs were chrome yellow to deep chrome. but sometimes there was a pinkish vinaceous spot in the center of the throat. Pueblo Rico. The sides may be black or light and are sometimes spotted. while the centers of the ocelli on head and sides were dull gallstone yellow. La Granja. wittei or D. In other examples the centers of the large ocelli on top of head and on sides. 54186. some of the frogs were almost uniformly black. CNHM Choc6: CNHM Andagoya. The light area on the legs was pinkish buff. the ocelli quite dark in contrast. t. so we can hope for comparisons of still larger series from future collections. The belly was solid black. La Selva. CNHM 54184. with chestnut to chocolate-brown or seal-brown squarish spots and. There are no outstanding differences in critical measurements D. Fortunately. chocoensis and a somewhat shorter one than D. t. the original yellow ocelli being a dull dark gray. histrionicus from the other subspecies of D. t. were received freshly dead at the National Museum on August 7. The few light areas on the belly were raw sienna. the spots being brilliant cadmium yellow. 14030. longitudinal stripes. 54227. histrionicus and D. 54234-47. 54211. appear to be intermediate between D. 54197. at 400 meters (CNHM 54229 and 54248-74). or they may have a branch of the light middorsal spot extending across the axilla. Specimens from Playa de Oro. Usually the back was ochraceous to ochre yellow. the glands above the groin were bright chrome yellow.) The belly was black except for the maize-yellow to dull-cream edge of the lower jaw. 139778. although it has a longer tibia on the average than D. 54190. . Rio San Juan. NATIONAL MUSEUM BULLETIN 2 88 spot on the tip of the chin and a median one on the posterior part of the abdomen. individuals of this group usually are abundant wherever they occur. 54222. 54188. A year after preservation. 81853 MLS (juv. and their colors were noted before they USNM were preserved. Caldas: Santa Cecilia. to separate Specimens Examined COLOMBIA AMNH Antioquia: Andes. 1962. Rio San Juan. 54206. 54231-2. from Quibd6. 54201-2. conjluens. arms.) . 54228. Choc6. occasionally. Bahia Utria. (In one example. tinctorius found in Colombia.30 U. 54208-9. 147164-203) from Playa de The lot of 40 examples (now Oro. t. 54224-6. 54213-20. south of Andagoya. t. Other frogs were pale olive-buff or drab gray. Finca USNM 75 mi.

the disk of which covers about one-third the tympanum. 145a. except for a few pustules on the sides and above the shoulders. lower Rio Atrato. metacarpal tubercles only slightly developed. its posterior border free and unnotched. — locality. west Playa de Oro. Can thus rostralis rounded. MLS 145. a small swollen area in place of a definite thumb pad. tributary of the 1913. — Boulenger. 1028. USNM 144945. a small oval inner metatarsal tubercle and a smaller round outer one. pi. Rio Sucio. 10-15 km. Body moderately elongate. tips of second and fourth fingers extending to base of penultimate phalanx of third. its diameter equal to its distance from tip of snout. about 1% times the interval between nostrils. USNM 147164-203. interorbital diameter twice that of upper eyelid. the upper jaw extending well beyond the lower. USNM 144947.12. when hind legs are bent at right angles to body. 27. ovate. When hind leg is ad pressed. no skinfold across the chest. third toe longer than fifth. 5. a small palmar callus. Fingers moderate. no glandular ridge behind eye. Rio San Juan. of BM Dendrobates tinctorius chocoensis (Posada Arango) Plate 4a-c 1869. heels slightly overlap. a series of small skinfolds on knee and heel. Choco. flat and sloping backwards in profile. figs. a very faint tarsal gland. a few flat granules behind the vent and on lower femur. Colombia). venter smooth. 6. Laurent. ending midway to heel. Skin of upper parts smooth. its greatest diameter about two-thirds that of eye. Pacific coast. 150626-9. vertical. heel reaches to posterior border of tympanum. an adult from three miles east of Andagoya. 1872. in postaxillary region narrower than greatest width of head. prominent. Choc6 Colombia. Dendrobates tinctorius chocoensis. Eye large. knee and elbow touch. USNM 124264. Upper Condoto. first finger a trifle shorter than second and fourth. its disk reaching halfway on antepenultimate phalanx of fourth toe. Phyllobates chocoensis Posada Arango (type Deyrolle. Tongue about one-third as wide as mouth opening. 104 — Description. no external vocal sacs in the male. USNM 144946. pi. free. snout broadly rounded at the tip when viewed from above. long.90-3. and in line with the upper lip. p. Toes rather short. Tympanum rather indistinct posteriorly. 12. p.— FROGS OF COLOMBIA — COCHRAN AND GOIN 31 upper Rio Atrato. p. a ventral disk. . separated from eye by an interval equal to one-fourth its large disks but no own diameter. 1913. Nostrils inferolateral. scarcely projecting. with lateral ridges.11. their distance from end of snout about two-thirds their distance from eye. on the road to Opogod6. 1942. MLS 227. Quibdo. Rio San Juan. when limbs are laid along the sides. the disk of which covers one-third the tympanum and equals that of third finger. loreal region slightly concave. 465. 1913.

.11. its posterior border free . tibia.5 mm... 1915. The dark dorsal spots are different in size and arrangement in all specimens.23-4. Dorsum olive-buff. Colombia).21. 15 mm. road to Opogod6.10.21. 11. The varied dorsal pattern of D. with an irregular sepia crossband on femur. but it can be easily distinguished from D. 1915. foot. an indistinct sepia band across the chest. spots on the back. while the belly is black.25. states A note with specimen color was "above merging shades of orange and yellow.10. The arms and legs often are light above. Pefia Lisa.t.. oval.78-80. but most of the Pen a Lisa and Rio Condoto examples have dark spots or bands on the limbs. Usually the snout is light.32 U. chocoensis because of its dark dorsum with small evenly — BM — scattered light spots. 75 (tjr pe locality. Condoto. vrittei also has a fairly short tibia. Tongue one-third as wide as mouth opening.21.— 13250). hand. from Andagoya.21. with a pale drab spot covering outer and lower part of knee.5.51-7. 10. D. 144941-4.82-78.10. Rio Condoto. 146S39. and two or three fainter ones on outer part of foot and across upper arm and forearm. from Andagoya.t. Rio San Juan. 12 mm. 35. 124262-4.. Head and body. Narifio. the borders of the lower lip fading to drab. 8. This subspecies tends to have a proportionately shorter tibia and hand than the other Colombian forms in the tinctorius group. p. 13 mm. NATIONAL MUSEUM BULLETIN 288 — Dimensions." below Remarks. 1916.52. t. Colombia. about 600 m. 1913. although it may have a dark spot in the center or on one side of it. their disks and tubercles fading to wood brown. femur.S. USNM Dendrobates tinctorius confluens Funkhouscr Plate 4d-f 1956. top and sides of head and snout immaculate. with several irregular sepia Color in alcohol.. palms of hands and soles of feet dark sepia.. Usually the sides and sometimes the chest are light. Specimens Examined COLOMBIA Choc6: BM CNHM USNM BM BM Andagoya.8181850-2. lower limb surfaces clove brown to sepia. head length. another on distal part of tibia. Ciudad de Madrigar. Nariflo. 1915. chocoensis suggests a "clown suit" of very irregular dark spots on a light ground. 10 mm. upper limb surfaces olive-buff.5 mm. Venter clove brown. and 1914. USNM SUNHM La 145800 (formerly Description.12. that in life the dark to black brown.4.5 mm. a paratype from La Ciudad de Madrigar. Dendrobates histrionicus confluens Funkhouser. head width. 145792. three mi.

tip of snout. 8. scarcely projecting. a pair of a slight skinfold across the chest.5 mm. ending at heel. head length. 13 mm. loreal region slightly concave. femur. are laid along legs are bent smooth (pustular under the lens). Toes moderate. nostrils. width. and chest covered by an Color in alcohol. foot. except for some small tubercles on femur near anus. When hind leg is adpressed. third toe much longer than fifth.. with lateral ridges. with minute black dots which merge to form black patches in axilla and around shoulder and in groin. as it is pale geranium pink above and on the sides and small slatecolored spots are scattered over the back. — Dorsum olive-gray with many small irregular black spots scattered over head. the upper jaw extending well beyond the lower. the edge of upper lip black. lower surfaces of legs. and tibia.. Canthus rostralis rounded but distinct. Nostrils lateral. disks and tubercles on feet and hands pale drab-gray. tibia. the disk of which covers half the tympanum. a heavy skinfold on heel and knee. 12. disk of fourth toe covering one-third the tympanum and a little smaller than that of third finger. posterior femur olive-gray with fine black dots. Fingers long.. Eye large. when limbs when hind the side. continuous with the upper lip. femur. body. 10 mm. and hands black. metacarpal tubercles rather weak. a slight tarsal ridge.5 — Head and body. vertical. Dimensions. separated from eye by an interval equal to one-fifth its own diameter. its disk reaching halfway on antepenultimate phalanx of fourth. Body elongate. heels overlap. ending in front of shoulder. 32 mm..FROGS OF COLOMBIA — COCHRAN AND GOLN 33 and unnotched. interorbital diameter equal to that of upper eyelid and equal to interval between nostrils. hand. a weak oval thumb pad present. 9. throat. weak ventral disk. their distance from end of snout one-half their distance from eye. heel reaches posterior edge of tympanum. a larger round palmar callus. Skin of upper parts corner of eye curving behind tympanum. truncate and sloping backwards in profile. a weak oval inner metatarsal tubercle and a larger round outer one.. The venter is smoke-gray.5 mm. chin. 13267 has retained some of its original color. venter smooth. feet. Tympanum distinct. venter slate color marbled with black.. free. 11. prominent. fourth longer than either and reaching to penultimate phalanx of third. head mm. and canthus with irregular black reticulations. knee and elbow slightly overlap. snout broadly rounded when viewed from above. a weak glandular ridge from posterior at right angles to body. sides olive-gray SUNHM . side of head olive-gray. its diameter equal to its distance from tip of snout. a lateral external vocal sacs in the male. in postaxillary region equal to greatest width of head. irregular black spot.5 mm. its greatest diameter about two-fifths that of eye. first finger much shorter than second.

61774. —Specimens Guayacana. Fingers and toes dilated into regular disks subdivided above. a large black spot occurs on center of throat. . Outer metatarsals united. USNM 147401-29. (paratypes). 1. in the are said series ANSP by Funkhouser 25349-52. 61775) from La Guayacana have been similarly identified by the present authors. The most noticeable structural difference and head the former.. Pichincha Province. The feet and hands are black above and below except for the disks and tubercles. The tip of the snout. preceded by a vertical black spot. A narrow black ring extends around base of arm and leg. In addition to these specimens mentioned by Funkhouser. Terminal phalanges T-shaped. t. Rio Mira. Specimens Examined COLOMBIA CNHM Popaydn.S. and nostrils are irregularly blackspotted. Remarks. Caxjca: NARitfo: Genus Phyllobates Dumeril and Bibron 1841. The femur and tibia are black below. P. bicolor Dumeril and Bibron). USNM 147317-9. p. which are smoke-gray. Vomerine teeth absent. Imbili. maxillary teeth present. Those spots that have faded to gray have the appearance of lichens. sternum a cartilaginous plate. and there are some fine scattered short black lines on the belly. the black area having an irregular margin. t. Phyllobates Durueril and Bibron.34 U. canthus. La Ciudad de Madrigar. free behind. Generic diagnosis. three others (CNHM 61772-3. An irregular black crescentic mark occurs just above the arm and some distance behind the tympanum. Tongue elliptic or subcordi- form. Several other frogs from the same place show some pink color. as the small irregular black marks occur at random over the upper surface. p. and merging on the femur with the black of the lower surface. Rio Mataje. sometimes weak. Narifio. (1956. in northwestern Ecuador) but to be closer to the former. NATIONAL MUSEUM BULLETIN 2 88 with the lower jaw rimmed with black. USNM 145800-1 [formerly SUNHM 13250. SUNHM 13151 (holotype). 13267]. 13251-66.700 m. Omosternum cartilaginous. from La to be 75) intermediate between confluens and sylvaticus (from nine kilometers west of Santo Domingo de los Coloradas. whose between D. widening in axilla and groin. confluens D. 637 (type species. The upper lip has a spot of bright geranium pink below the eye. —Pupil horizontal. tinctorius is the significantly longer foot of width and tibial length also appear to be slightly greater.

thorntoni and nostril. "Cuba" in error. size to 42. size to 27. subpunctatus subpunctatus d2 Toes one-fourth to two-thirds webbed. and reticulations. belly immaculate or finely spotted. 1 . mm Phyllobates bicolor Bibron Plate 6a-c 1841. pi. heel reaching between anterior and posterior corners of eye. mm P. 1 . 1 P.FROGS OF COLOMBIA Key Toes a'. Dumeril and suggested as probably Colombia by Dunn. d Heel to anterior eye. heel reaching between posterior border of eye and nostril. toes free. mm .. or subequal. . d Toes webbed at base. toes one-half webbed. First finger longer c1 . P. third size to 25. and second . sometimes mm P. subpunctatus walesi Toes one-third to one-half webbed.5 A b2. e1 . toes webbed at base to one-eighth webbed. 29 (type locality. heel reaching between posterior border of tympanum and eye. P. size to 22 mm. boulengeri Chin and anterior part of belly black. skin resistant to abrasion. toes not webbed. heel to between eye f 2 . P. size to 22. size to 22 mm. posterior part marbled dark and light. anterior corner of eye. light lateral or dorsolateral stripes.5 e3. mm b2 . d d2 Belly with coarse dark reticulations continuing on throat .5 P. Belly spotted. size to 26 . femoralis subequal. heel to center of eye. brunneus d2 Heel to center of eye. than second. P. size to 18 P. chocoensis Belly immaculate. c1 . heel to tip of snout. latinasus d2 . belly pinkish buff with small scattered sepia spots. mertensi Toes webbed at base to two-thirds webbed. vergeli fingers subequal. b free. bicolor faint. pratti e2 Belly immaculate except for a dark line around lower jaw and a patch of dark dots below shoulder. mm . 1944c). size to 32. . size to 22.. Toes nearly one-fourth webbed. belly drab with a few darker spots below femur and tibia. e Belly immaculate buff. c2 . size to 36. c2.5 First d mm . belly drab with heavy dark reticulations. p. bl . 235. a pair of short parallel stripes on chin. and second First fingers mm a2 . mm l . first finger slightly longer than second.5 .. heel to Belly with dark spots /'. — . Toes one-fourth webbed.5 and fourth toes with a slight web.. 1 . size to 28. P. toes one-third webbed. mm c2.5 P. c First finger shorter than second. P. First finger shorter than second. belly usually slate color posteriorly (in alcohol). heel to posterior border of tympanum. other toes free. . P. Phyllobates bicolor Bibron.5 mm Belly with a coarse dark network. inguinalis Toes webbed at base to one-eighth webbed. . belly immaculate drab. lighter anteriorly. light dorsolateral stripe. and forming . — COCHRAN AND GOIN 35 Colombian Species of Phyllobates to barely webbed. or No x .5 P. size to 30. . l . size to 25 mm. palmatus First finger longer than second.

p. heels touch. — Dimensions. Description. upper limb surfaces with numerous.5 mm. Toes free. metacarpal tubercles well developed. tongue half as wide as mouth opening. 14 mm. free. its disk reaching halfway on antepenultimate phalanx of fourth. when limbs are laid along the sides. 1961. pi. 491. 18. p. the tips of first and fourth reaching to base of penultimate phalanx of third. Tympanum very distinct anteriorly. the disk of which covers about one-fourth the tympanum. ovate. a slight tarsal ridge ending in a heavy diagonal gland midway to heel a heavy skinfold on heel and knee. 107. without lateral ridges. 1963. 77. Eye large.] Colombia). when hind legs are . interorbital diameter one-half that of upper eyelid. 1841. disk of fourth toe covering one.] Colombia). Phyllobates Popayan. Phyllobates melanorrhinus Berthold.. Head rather short.third the tympanum and slightly larger than that of third finger. a very slight skinfold across chest. a larger palmar callus.— Cochran." [Antioquia. 1. its diameter equal to its distance from tip of snout. femur. fig. 638— Giinther.. p. 12 mm. less so posteriorly. 1923. heel reaches center of eye. Cauca. its posterior border free and unnotched. projecting. 7 (type locality. a small oval inner metatarsal tubercle and a still-smaller. Nostrils lateral..5 mm. vertical. with faint lateral ridges. loreal region flat. 25. 42. NATIONAL MUSEUM BULLETIN 288 1847. nicefori Noble. 16 mm.36 U. p. head width. Bibron. truncate in profile. first finger a little longer than second and equal to fourth. knee and elbow overlap. no external vocal sacs apparent in the male. — USNM 137360. very small slate-gray dots which are set closer toColor in alcohol. no glandular ridge from posterior corner of eye. an adult female from Popay&n. 20 mm. Body moderately elongate. tibia. head length. Canthus rostralis rounded. except for a few small granules on posterior part of abdomen and on lower femur beside the vent.. 1 (type locality. foot.—Gorham. third toe longer than fifth.. 1944c. — Dorsum uniform pearl gray. Fingers moderate in length. When hind leg is adpressed. round outer one. the upper jaw extending well beyond the lower. fig. no ventral disk. its greatest diameter twothirds that of eye. "town of Andes. 14. p. bent at right angles to body. with distinct disks. Skin of upper parts smooth (finely pustular under the lens) and very resistant to abrasion. their distance from end of snout one-half their distance from eye. .— Dunn. p. prominent. separated from eye by an interval equal to onefourth its own diameter. Colombia. 1957. 13. equal to interval between nostrils. in line with the vertical upper lip.. snout broadly rounded when viewed from above.S. venter smooth. 1858. in postaxillary region narrower than greatest width of head. hand. Head and body. [Cauca. p. 39. 90. a small round thumb pad present.5 mm. p.

54221. BM 1910. USNM 14506. lower Rio Calima. USNM 139776-7. The toes lack webs. tibia. 5. In alcohol. 54207. 13540.6 to species — that more than a normal coefficient of variability is.) this frog and 11 others from the same region had bright redorange backs.7. ANSP 25490. west of Santa Cecilia.. 149723-7. and the upper surfaces of the femur. Cauca: Choc6: 53168. USNM 146630-1. 3. 12 km.0.FROGS OF COLOMBIA —COCHRAN AND GOIN 37 gether on outer parts and merge into a nearly uniform slate color on lower surfaces of limbs and posterior part of belly. and disks and tubercles of hands and feet dark brown. Pueblo Rico. and anterior half immaculate. USNM Carton de Colombia. soles and palms slate gray.200 m. a black crescentic nostrils and borders black. lower surfaces of limbs black. MedeMn. AMNH 54183. 39081. 54233. D. The body among the largest length of 42. this shows no from 4. Atrato River.11. The skin is most of the recently collected specimens the older examples are quite free of holes or tears. Rio San Juan. 10-15 km. a triangular black mark on center of belly. the Peruvian example (CAS 85151) is straw yellow to color. USNM 144994. Specimens Examined COLOMBIA Antioquia: Andes. throat. critical its 6. from Buenaventura.53. and many of as heel reaches the anterior border of the eye or as far as the nostril in over 80 percent of the frogs examined. cream —In Remarks. USNM 53170-1. south of Andagoya.. chest and throat with scattered black dots. 337-262—70 i . 54212.C. in less than 20 percent it reaches the center or the posterior border of the eye. mark around lower of lips much anterior rim of tympanum. Rio Raposo. Juntas de Tamana. and arm were yellowish. 54196. Quintana. 54204-5.21. The measurements. chest. the disks and tubercles of belly cinerous. USNM 137359-70. Virology Field Station. 54189. Rio Truando. USNM 48865. El Pailon. 122104 [formerly USNM 4349]. ANSP 25755. 147204-7. While alive (at the National Zoological Park in Washington. 75 mi. 54187. CNHM 54183. USNM 145105. paler.000 BM 1915. USNM 147225-6.5 millimeters places Phyllobates bicolor species of the genus. 150632-7. mountains above USNM 150638-42. AMNH Caldas: AMNH 14028-9 (paratypes of P. Rio Sipi headwaters. PERU: Tingo Maria. River. and the tongue is unnotched. ft. upper Atrato Playa de Oro. with large black spots between the shoulders merging into a black network on posterior half of back. CAS 85151.46-7. resistant to abrasion. Valle: ANSP Camp 25756. nicefori Noble). Popayan. 54193-4. lower Calima River. Sierra de Baud6. USNM 151381.10. Playa de Oro.

1963. in postaxillary region slightly less than greatest head width. a weak tarsal ridge. Prosthcrapis boulengeri Barbour.. 1926. Width of tongue about one-half that of mouth opening. disk of third finger covering about a deep furrow along the edge of the it. head length. snout rather short. from Gorgona Island. disk of fourth toe covering one-fourth the tympanum. broadly truncate at the tip when seen from above. not ridged laterally. p.S. Fingers long. lip. 89 (type locality. with narrow lateral ridges. 8. 101 (type 1909. 1883) Barbour. extending beyond base of ante- penultimate phalanx of fourth. MCZ USNM 52406 [formerly Description.5 mm. p. nostrils lateral and well below the canthal angle. the upper jaw extending well beyond lower. 5. 1926. spatulate. a small inner and a smaller outer metatarsal tubercle. 25. — Parker. Canthus rostralis rounded but distinct. foot. 8. Tympanum distinct. Color in alcohol.5 mm. when limbs are laid along the sides. Eye large and prominent. venter with a few granules beneath anus. head width. cotype of an adult female with eggs. 19. ending in a diagonal gland across the tarsus proximal to the metatarsal tubercles.. separated from each other by an interval of about 1% times their distance from eye. a small palmar tubercle. about one-half the diameter of eye and located immediately behind it.5 mm. hand. third toe much longer than fifth.. Prostheraspis femoralis (not of Boulenger.5 mm. but narrower than distance between the nostrils. NATIONAL MUSEUM BULLETIN 2 88 Phyllobates boulengeri (Barbour) Plate 7a-p 1905. heels fail to touch. Barbour and Loveridge. when hind legs are ad pressed.. with boulengeri]. 38 U. rounded in profile. 6. with a very minute indentation on its free posterior border. their distance from end of snout slightly over half their distance from eye. first finger a little longer than second. tibia. Head and body.— Gorham. 7 mm. and a pair of small flaps above it. loreal region flat. 8. subarticular tubercles of toes smaller and less evident than those of fingers. Bleached to a nearly uniform pale ecru drab. Dimensions.. but with traces of a sepia stripe on the side of the head from nostril — — . Body moderately elongate. Phyllobates boulengeri. interorbital diameter about 1% times the width of upper eyelid. free. 323. one-fourth the tympanum. its subarticular tubercles of fingers large. heel reaches to posterior corner eye. 1929.5 mm. locality. toes not webbed.— .. p.5 mm. 553. p. part. which flares out a little below diameter equal to its distance from tip of snout. Gorgona Island. Skin not easily torn. femur. Colombia).— 2422. knees and elbows slightly overlap when hind legs are laid at right angles to body. Gorgona Island). Colombia. smooth (minutely pustular) above and below. p.

Except for usual variations in darker or lighter back and more or fewer dark reticulations on lower surfaces. posterior femur clove brown with numerous very small light dots. sepia dorsoa above a narrow. Phyllabates boulengeri deserves to be kept distinct from Phyllabates jemoralis (Boulenger) because of small but constant differences. the disks and tubercles lighter. white line is the — the first fingers being a little longer. tibia and foot with three irregular ocellated sepia crossbands. The wide dark lateral band bordered above by the narrow straight lateral stripe same in all the specimens. white stripe going straight back from posterior corner of upper eyelid to upper insertion of femur. The fingers are long and slender. A large series of specimens (USNM 145248-300) was secured by Dr. around armpit. soles of feet and palms of hands drab gray. as a rule. but in jemoralis a narrow white stripe. a white line from near snout tip along upper lip to shoulder. thicker. and part way along upper arm. bordered below by a dark stripe. the other specimens of this series have a color pattern very similar to that of the described individual (USNM 145268). side of head and body with a wide clove brown stripe extending to groin. Some have the lateral halves of the throat dark. with an irregular white median line and white lip borders. another stripe on side of body from armpit almost to and two paler brown stripes on the posterior femur. snout paler drab-gray faintly marbled with sepia. Following is a description of to groin. than the second. For instance. In the Gorgona Island series. the color pattern of an individual material: lines on Dorsum drab-gray with either side of the midline. venter white with coarse clove brown reticulations continuing on throat and forming a pair of short parallel stripes on the chin. The throat of boulengeri shows a wide . anterior femur white proximally with irregular clove brown longitudinal markings. sharp. is found along the lateroventral region paralleling the wide dark lateral stripe. arm with small sepia spots. (USNM 145268) in this fresher row of irregular sinuous sepia and a darker. the wide dark lateral stripe meets the white belly color directly in boulengeri. Medem on Gorgona Island in 1961. which rarely extends as much as one-eighth the distance along the fourth toe. the toes usually show a minute trace of a web. Remarks. The heel reaches either to the anterior border or the center of the eye in 95 percent of the frogs measured and to its posterior border in the others. the femur with several diagonal clove brown stripes separated by narrow white lines. Center of throat also with pale brown markings.FROGS OF COLOMBIA — COCHRAN AND GOIN 39 tympanum. upper limb surfaces drabgray. can thus rostralis and tip of snout with a light ecru drab line. lower surfaces of limbs white with scattered clove brown spots. its lower edge irregular. The largest frog is 22 millimeters in head-and-body length.

1935. 1957. 149723-7. loreal region flat. Huallaga River. p. separated from each other by an interval about 1% times their distance from eye. Caqueta. 1952. In both species the skin unduly subject is not to abrasion. p. and the dorsum and venter of the former species are while the back is smooth (minutely pustular under the lens). nostrils lateral. 1957. p. 1S84. fig. Peracca. their distance from end of snout about one-half their distance from eye. Narino: Imbilf. rounded in profile. while the posterior belly is light with black spots. Barbour and Noble. Gorham. 138 (part). 78. p. the upper jaw extending well beyond the lower. 1904.— Goin and Layne.) —Dunn.— Dunn. 401. 1882. p. Canthus rostralis rounded but distinct. Colombia. USNM USNM Phyllobates femoralis (Boulenger) Plate 8a-c — Boulenger. 1919. 14731-50. Pena 1920. Phyllobates inguinalis (not of Cope). snout moderate.— Beebe.— Crawford.— 40 U. p. pi. 18S4a. The foot of boulengeri is signifi- cantly shorter than that of femoralis. p. its free posterior border with three weak lobes. spatulate. a species associated solely with the mainland. 146872. Scarcely a tear appears in the skins of the recently collected series of examples from Gorgona. 15. 118. — . 145248-300. 9. Prostherapis inguinalis (not of Cope). Lisa. — USNM — p. 36. p. 118232-3 cotypes of femoralis Barbour). p. p. placed below the canthal angle. 78. 1920.S. at 300 ft. Width of tongue more than half that of mouth opening. 1898a. lower Calima River. Dendrobales aurotaenia Boulenger. 1 (type locality. 25. Specimens Examined COLOMBIA USNM Cauca: Gorgona Island: 52406 (cotype of boulengeri Barbour). 1958. 1957. distinctly granular in It was interesting to find that specimens of Phyllabates boulengeri from Narino and Valle are identical to those from Gorgona Island instead of being more similar to Phyllabates femoralis. 1029. 1957. p. broadly truncate at the tip when seen from above. 1963. 635 (type locality. Parker. and its chest and belly are white with some dark spots and reticulations. Phyllobates femoralis. p. Valle: Camp Carton de Colombia. In femoralis the throat and chest and anterior half of belly are uniformly black. 209. — Dunn. Lutz and Kloss. 412. with a narrow furrow along its lower boundary from nostril to below the eye. NATIONAL MUSEUM BULLETIN 288 black sinuous stripe on each side. 102. p. Condoto. most examples of femoralis. 1926b. [Choc6] Colombia. 1882a. — — Phyllobates auro-taenia. p. p. Description. northern Peru) 1886b. Yurimaguas. Prostherapis femoralis Boulenger. 1935. 506. 77. 671. Noble. 1913. . an adult from Florencia. p. 1931. vertical. Rfo Mira. 104.

— 6. 9 mm. short. No inguinal or lumbar gland. belly and lower leg surfaces white with many large black spots. subarticular tubercles of toes smaller and less evident than upper Eye lip large . below tympanum. 11. first finger longer than second. heel reaches to center of eye. skin of arms nearly smooth. about one-half the diameter of eye and located immediately behind eye. very irregular in outline. head width. knee and elbow greatly overlap when hind legs are bent at right angles to body. venter slate-gray anteriorly. and prominent. A white line beginning below nostril andrunnin g along edge of upper lip and to the shoulder ends in a large crescentic white patch. and straight backward along side. third toe longer than fifth. a narrow white lateral line from below arm along side to lower anterior part of groin. venter smooth. Tympanum distinct... a large squarish white spot in upper part of groin and another at armpit. with a narrow white dorsolateral above eye. hand. a third white line. its diameter equal to its distance from end of snout.. from axilla to groin along lateroventral border and paralleling the two upper lines.. disk of fourth toe covering almost onehalf the tympanum. tibia. 8 mm. free. the other toes without webs. heels slightly overlap. Dimensions. that of upper leg surfaces coarsely granular. with small disks. Color in alcohol. transverse glandular ridge across tarsus about one-third the way from inner metatarsal tubercle to heel (which suggests the tarsal tubercle found in some species of Physalaemus) a small but distinct inner metatarsal tubercle and a smaller outer one. finely granular on tip of snout and on side of body. coarsely granular on back. legs barred with lighter slate gray diagonal bars. Body rather elongate.5 mm. 10. third and fourth toes with a very short web between them. Skin not easily torn.5 mm. width of head when hind legs are ad pressed. head length. 12 mm. subarticular tubercles of fingers very large. —Dorsum black. a heavy. reaching halfway on antepenultimate phalanx of fourth. in postaxillary region equal to greatest . the posterior half of line from above nostril. palms of hands and soles of feet spotted heavily with black.FROGS OF COLOMBIA —COCHRAN AND GOLN 41 rounding outwards very slightly beyond the loreal region. Head and body. femur. those of fingers.. their corresponding upper surfaces white with a few black dots. when limbs are laid along the sides. . interorbital diameter a little greater than that of the rather broad upper eyelid. without lateral ridges. above arm insertion. Fingers moderately long. subarticular tubercles of toes very small. 25 mm. a large palmar tubercle present. throat and chest black. equal to the distance between nostrils. continuing on edge of upper lip. . fourth rather short. and straight back to leg insertion.5 mm. disk of third finger covering about one-eighth the tympanum. a second white line beginning below nostril. foot..

CNHM MCZ Putumayo: UF 33275-7 (tadpoles). — Diagnosis. chest. MLS Carlo Guapaya. Umbria. USNM Valle: USNM USNM USNM Phyllobates mertensi Cochran and Goin Plate 6g-i 1964. USNM CNHM 81810. 255 (type locality.6 in the series of 16 examples examined indicates that the series is too small for an adequate estimate of variation in the critical measurements. dark gray or black throat. and anterior belly at least in the males and the heavily reticulated surfaces of the posterior belly and lower leg. The coefficient of variability in the head width and hand length 9. — — Specimen8 Examined COLOMBIA Amazonas: Puerto Narifio. Cope does not mention such spots in his original description. 199. 506). 127210-3. USNM Leticia. 147268. PERU: Ceja de Montana. Phyllobates mertensi Cochran and Goin. 151382. Caldas: Villa Maria. Huanuco. belly NATIONAL MUSEUM BULLETIN 2 88 and lower limb surfaces marbled with coarse black and white spots. — — . Resembles Phyllobates chocoensis and P. 61790.S. Sierra de la Macarena.42 U. Mocoa. Remarks. heel reaches to the center or anterior corner of eye in about 95 percent of the specimens at hand. ANSP 25370-1. Quebrada Aqua Dulce. had been confused. —According to — dorsal surface Parker (1935. Morelia. Acacias. Buenavista. and an indistinct light diagonal stripe or traces of such a stripe across the side to the groin. p. inguinalis. Differs from these and from all other Colombian species of Phyllobates in having a very short tibia (between 35 and 42 percent of the total length) with the heel reaching a point between eye and posterior border of tympanum. Cauca). P. granular is p. 10016. The axillary and inguinal spots of femoralis are pink in life. 127202. 4a. femoralis is recognizable by the combination of a granular back and a peculiar ventral coloration. south of Villavicencio. and to the posterior corner of eye in the others. Caqtjeta: Meta: Florencia. separating with which it perceptible in the type specimen. Domo Santa Clara.2 and 10. 18. USNM 146S63-6. 146872-3. The back is very granular — — in most The of the specimens examined. bocagei in having a thickset body. broad snout. but some light areas in these regions are still femoralis. BRITISH GUIANA: Kartabo. MLS 4. a short. Quintana. the glandular — or a distinguishing feature of Phyllobates it readily from the smooth-backed P. Virology Field Station. Rio Raposo. 118075. near Popayan. JP 0408a-b. Orellana.

—Dorsum brown. venter smooth (finely pustular with magnification). separated from eye by an interval equal to three-quarter its own diameter. a small oval inner metatarsal tubercle and an outer one of about equal size. heel reaches posterior border of tympanum. tibia. . with a faint irregular darker stripe above the lip border. — Dimensions. knee and elbow just meet. Toes moderate in length. When hind leg is adpressed. an ecru drab crescent in front of arm insertion and in axilla. interorbital distance 1% times that of upper eyelid. Tympanum distinct.. first finger equal to second. its diameter 1% times its distance from tip of snout. in postaxillary region equal to greatest width of head. 12.. a small oval thumb pad present. prominent. edge of lip slightly paler than side of head. scarcely projecting. free. when limbs are laid along the side. tympanum fawn color. the upper jaw extending well beyond the lower. side of head seal brown. webbed at the base. 54456 (holotype). and with traces of dark bars across upper surfaces of thigh and shank. metacarpal tubercles well developed. sloping outward to the upper lip. its posterior border free and slightly notched. a faint ventral disk. a large round palmar callus. oval.5 mm. hand.. its disk reaching to center of antepenultimate phalanx of fourth. a little greater than interval between nostrils. its greatest diameter one-half that of eye. fourth longer than either and reaching to base of penultimate phalanx of third. 10. near Popayan. 30. head width. third toe much longer than fifth. the disk of which covers about one-eighth the tympanum. Tongue half as wide as mouth opening. a pair of lateral external vocal sacs in the male. a faint tarsal ridge ending diagonally across tarsus midway to heel. this area bordered by darker brown and sharply set off from the paler fawn color of the lower surface of the arm. with lateral ridges.5 mm. foot. heels are widely seperated.5 mm. disk of fourth toe about one-third the tym panum and larger than that of third finger. a heavy skinfold on heel and knee.. femur.5 mm. Eye large. irregular darker spots seal on each side .5 mm. 9 mm. 7. Nostrils lateral. Color in alcohol. truncate in profile. 11. an adult female from Quintana. fading out at midbody but again perceptible near the groin. upper arm chocolate above. Cauca. Can thus rostralis bluntly rounded. with some very indistinct of midline. Head and body. Colombia.. smooth (thickly pustular under the lens) a small parotoid swelling extending backwards as a narrow lateral gland.5 mm. side seal brown.. when hind legs are bent at right angles to body. head length. their distance from end of snout three-quarters their distance from eye. 11. no skinfold across the chest. loreal region faintly concave. Skin of upper parts resistant to abrasion. snout broadly rounded when viewed from above. Body stout.FROGS OF COLOMBIA — COCHRAN AND GOIN 43 — CNHM Description. Fingers fairly long.

Specimens Examined COLOMBIA CNHM 54456 (holotype). opening. D. palms of arm immaculate ecru drab. sides seal brown in alcohol. Toes one-third webbed. the upper jaw extending well beyond the lower. Antioquia. hands and soles of feet seal and inner surfaces seal brown. sloping down rostralis distinct. lower thigh with a pale drab area on proximal half and with fine seal brown reticulations over the remainder. Can thus loreal region vertical Eye and nearly flat. size to 32. and CNHM USNM 147228-9 (all paratypes). snout nearly truncate when viewed from above. its diameter 1% times its distance from tip of snout. Antioquia. Antioquia. — Tongue a little more than half as wide as Description of holotype. their distance from end of snout about one-half their distance from eye. with a small light drab spot but no definite light line. 163a-b. near PopaUSNM 151180-2. interorbital diameter equal to that of upper eyelid. an adult female from Medellin. 54341. USNM 147512. 818778. 38806-7. 54428. 14020. Johnson. Purace. —AMNH 1347. skin resistant to abrasion. outer limb brown. chocoensis. slightly projecting. aDd belly. 44042-3. 1375. and webs ecru drab. 1363. all from Medellin. without any light dorsolateral or diagonal stripes. 1370. Tympanum distinct. tongue and other features. yan. foot. 44041. oval. It. Nostrils lateral. O. the lateroventral border very irregular. adpressed heel usually reaching to between eye and nostril. near Popayan. a little less than interval between nostrils. tubercles. 63889-90. belly with dark spots and reticulations on a light ground. to the upper lip. occasionally to posterior border — of eye. a few pale spots near the in front of groin swollen glandular area just above the groin. from Andes. MLS 163. 44045-6. the disks.5 The absence of any mm. collected by Paratypes—AMNH. Antioquia. broadly rounded in profile. light dorsolateral stripe or light diagonal stripe crossing the dark area in front of the groin places the new species close and P. chest. all from Yarumal. Quintana.S. venter ecru drab with a coarse seal brown reticulation over chin. It is distinguished from bicolor and from both bicolor and chocoensis by its notched to Phyllobates bicolor by its webbed toes. its posterior border free and very slightly mouth notched. separated from eye by an inlarge. lower surfaces of shank. Colombia. Diagnosis. USNM 147511. its greatest diameter about one-third that of eye. prominent. USNM 147227. all AMNH CNHM 14016. Cauca: CNHM Phyllobates tliorntoni. . new species Plate 8d-f Holotype.44 NATIONAL MUSEUM BULLETIN 288 XJ.

first finger shorter than second. sides seal brown. hand. Often the chin and chest are spotted. 7 mm. venter smooth. 9 mm. rather short.1 to 46. the lens is The belly appears it smooth reaches to the posterior at first glance.5 millimeters in length. and forearm with seal brown crossbars. a round palmar callus. . brown. Remarks. a weak. When hind leg is adpressed.FROGS OF COLOMBIA —COCHRAN own terval equal to one-half its AND COIN 45 diameter. knee olive. a pronounced skinfold on heel and knee. seal lateral stripes. mm. mm. metacarpal tubercles scarcely developed. measures 32. side of head and body also seal brown. in which corner of the eye. — Dorsum Color in alcohol. with large disks. no external vocal sacs in the male. knee — mm. soles of feet and palms of hands tawny-olive. the disk of which covers about one. with lateral ridges. its disk fairly large. disk of fourth toe covering nearly one-half the tympanum and larger than that of third finger. but seen to be pustular. when limbs and elbow overlap. 10 mm. both of these reaching to base of penultimate phalanx of third. 27 mm. third toe longer than fifth.7 to 49. free. becoming wood brown on upper lip where some pale spots of wood brown appear in the darker area. no skinfold across the chest. Skin of upper parts resistant to abrasion.. and sometimes three or four large light spots appear on the side of the belly well below the groin. a female containing eggs. tibia. upper femur. and in one specimen In adults the belly is it is under faintly granular. head length. foot. a small oval thumb pad present. tibia. Body elongate. No evidence of any well-defined black or white width. merging irregularly with the tawny-olive ventral color and lacking a light diagonal dorsolateral stripe. others of —This frog has a proportionately longer femur than any (with which genus Colombia except Phyllobates measurement overlaps very slightly— the range in in its the critical bicolor critical P. when hind legs are bent at right angles to body. Head and body. . belly with dark sepia spots and reticulations on the tawny-olive ground. heel reaches to center of eye . heels are separated. Dimensions.. The heel reaches between the center of eye and the nostril in all but one example. no glandular ridge above tympanum. and in bicolor from 46. 12 . which is equal to fourth. 12 . Toes one-third webbed. oval inner metatarsal tubercle and a smaller oval outer one. the dorsolateral region a slightly paler color. heavily spotted or reticulated.third the tympanum.8 percent). throat faintly marbled with olive. smooth (minutely pustular under the lens). . chest and lower femur tawny-olive with scarcely any dark spots.7 percent. head are laid along the sides. Fingers moderate. reaching to base of penultimate phalanx of fourth. in postaxillary region narrower than greatest width of head. thorntoni being from 46. no definite tarsal ridge. 9 femur. The largest example (AMNH 14016).

25. less so posteriorly.2. free. 1944c. Colombia). heel reaches anterior corner of eye. not easily torn. Hyloxalus chocoensis. oval. 1963. Antioquia. first finger shorter than second. p. separated from eye by an interval equal to one-half its own diameter. heels are separated. Colombia. equal to interval between nostrils. no glandular ridge behind eye. prominent. one-half webbed. an adult from south of Santa Barbara. venter pustular with a few fine glandular ridges on posterior half. When hind leg is adpressed. a very small oval thumb pad present. which covers nearly half the tympanum. Nostrils lateral. Body rather stout. its disk reaching to base of antepenultimate phalanx of fourth. 1963. Toes moderate. merging gradually with the upper lip. A weak bony ridges on top of head. rather blunt. Description. Prostherapis chocoensis. its posterior border free and unnotched. whose Colombian collections Phyllohates chocoensis (Boulenger) Plate 9a-b 1912. their distance from end of snout about one-half their distance from eye. in postaxillary region nearly equal to greatest width of head. — Dunn. skin normal. third toe a little longer than fifth. disk of fourth toe covering about half the tympanum and approximately equal to that of third finger. its diameter equal to its distance from tip of snout.S. with numerous irregular fine glandular ridges. Choc6. metacarpal tubercles well developed. —BM 1929. wide as mouth opening. a large heart-shaped palmar callus. when limbs are laid along the sides. NATIONAL MUSEUM BULLETIN 288 Wilmot A. loreal region vertical. with some elongate lateral tubercles on sides . It is appropriate to dedicate this species to Dr. interorbital diameter iy2 times that of upper eyelid. with lateral ridges and rather large disks. a pair of indistinct wide dorsolateral ridges. 190 (type locality. the upper jaw extending well beyond the lower. 1944.46 U. a distinct oval inner metatarsal tubercle and a smaller rounded outer one. Can thus rostralis curved. Hylixalus chocoensis Boulenger. Skin of upper parts finely pustular or shagreened. a distinct skinfold on heel and knee. truncate and sloping slightly backward in profile. Tongue almost two-thirds Tympanum pair of as distinct anteriorly. . 519. Rio San Juan. flat. Thornton. p. Eye large. have aided greatly in the present study. p. — Gorham.1. its greatest di- ameter about one-third that of eye. slightly projecting. snout nearly truncate when viewed from above. Fingers moderate. a sharp tarsal ridge. ending diagonally halfway to heel.5. knee and elbow overlap when hind legs are bent at right angles to body. fourth longer than either and reaching almost to base of disk of third.

4. the width and length of head as compared to length of body are greater in Phyllobates chocoensis than in P. and is continued in lessened brilliance or dorsolateral lines. 10.2.. of . a distinct ventral no external vocal sacs in the male. as the two appear very similar at first glance. throat and chest olive-buff.1. The dark dorsal areas here appear as two coarse chevron marks. subequal in bocagei. — Remarks. Head and body..2.5 mm. on top of the arm. with narrowed prolongations of the dark crossbars and indistinct gray reticulations.. 10 mm. head length. 2. otherwise the patterns both species are very similar. —Dorsum and lateral regions slate-gray. The heel reaches to the anterior eye in the four examples of chocoensis from Santa Barbara. femur.6). a pale gray axillary spot. which is 1% times the width of the upper eyelid in chocoensis and is equal to it in bocagei. with a large indistinct olive-gray spot behind each eye continuing aloDg the back almost to sacrum.FROGS OF COLOMBIA and below angle —COCHRAN AND GOIN 47 no skinfold across the chest. Antioquia. of jaw. apparently Dimensions. becoming much coarser on belly. and forming with its fellow an indistinct )( mark. their points nearly meeting. inguinalis or P.. foot. Webs between toes white. 27. disk.3) has only two small white spots on the upper lip.5 mm. mm. 10 mm. The large white axillary spot is very distinct. The snout of chocoensis seems wider. tops of legs and arms with a very wide slate-gray bar and one or two narrower and lighter ones on a pale olive-gray ground..1.5 mm. 8 tibia. According to the critical measurements. although it was said to reach the tip of ensis. lower surfaces of legs and arms with indistinct traces of a similar gray network. No light vertebral line is visible in the two adults or in the two immature frogs (BM 1929. Color in alcohol.. a fine gray network pattern on throat and chest.1. upper lip slate. lower surfaces of arms and legs deep ochraceousbuff. the last one below and behind the eye. however. lateral tubercles pale drab gray. while these two fingers are equal in in bocagei. The minute white spots on the upper lip of chocoensis seem to be absent in bocagei. 1929. 13. 13 mm. posterior femur dull ochraceous buff. — head width. The first finger is a trifle shorter than the second in chocoensis. and this fact is borne out by the interorbital diameter. thorntoni. with a narrow light edge and three or four small white spots near the margin. also the third toe is a little longer than the fifth in chocoof The degree webbing between the toes is a full half in chocoensis. hand. Boulenger was right in considering his "chocoensis" allied to bocagei Espada. while it seems less in bocagei and the margin of the web between the toes appears more deeply incised. No light vertebral The other adult at hand (BM 1929.

but it would not be surprising to find it in one of the southernmost states of Colombia. — Barbour and Noble. nearly vertical and slightly rounded in profile. littoral of the state of Rio de Janeiro. of Cuiaba. Peru). 1941. 1931. 30 mi. Phyllobates brunneus (Cope) Plate 6d-f 1887. Colombia. 166. 1957. Lutz and 673. 1964b.S. bocagei appears to have been recorded only from Ecuador. eastern Ecuador). Brazil). 1887a. 1961. p. 1955. p. chocoensis comes nearest to bicolor and thorntoni. [=Chapada]. — Ruthven. fig. fig." Apparently these two species are not closely related to P. Brazil). p. broadly rounded at the tip when seen from above. it is separated from bicolor by having its toes one-half webbed. with its easily torn skin.3-6. — USNM Description. an adult from Villavicencio. 1922. p. 1925.— Cochran. Phyllobates subpunctatus (not of Cope). 36. p. When grouped with species having an abrasion-resistant skin combined with the absence of light lines on the side and back. p. p. near Kaieteur Fall. So far. Phyllobates Boulenger. nostrils lateral. 1964a. preserving. 49. both chocoensis and bocagei appear to have abrasion-resistant skins that hold up well under the ordinary vicissitudes of collecting. northeast 1918. Gorham. palmatus. 146907. 1925. p. and shipping. 5. having about the same degree of webbing between the toes. 54 (type locality. 1959. p.— brunneus. p. 1922. 61. 311. and from thorntoni by having its tongue entire. 304. with a very slight indentation on its free posterior border.. Eupemphix olfersiodes A. p. Rio Pastaza. just . Meta. Specimens Examined COLOMBIA Antioquia: South of Santa Barbara. p. 2 (type locality. Prostherapis brunneus Cope. Lutz. BM 1929. Unlike palmatus. as well as by other in spite of the three characters. 138 (type locality. Width of tongue about three-fourths that of mouth opening. 48 U. although it has not been noticed heretofore. Kloss. 1920. Phyllobates intermedius Andersson. British Guiana). locality. 1923a. p. Roque.1. 1923. p. 280 (type locality. Villavicencio. 77— Gines. spatulate.— Dunn. snout quite short. the upper jaw extending well beyond the lower. 401. 34 (type locality. In the Ecuadorean bocagei the heel extends to "in front of eye. p. Phyllobates peruvianus Melin. 428 Mato (type — "Chupada" Grosso. The abrasion-resistance of skin appears to be a useful character for separating other web-toed species from palmatus. 1963. Dendrobates ranoides Colombia) 1920. 25. 1952. Hyloxalus bcebei Noble. NATIONAL MUSEUM BULLETIN 2 88 the snout in the unique type of the latter. p. projecting. p.2. 1945.— Rivero. 14. As noted elsewhere. 134.

width. fourth nearly as long. disk of fourth toe a little larger than that of third finger and equaling nearly one-half the diameter of the tympanum. but a slight longitudinal \vrinkling of the skin on either side of the throat below the jaw commissure. with only a slight transverse channel along its lower border. 9 mm.. free. not ridged. 7. when hind legs are bent at right angles to body. Can thus rostralis distinct and fairly sharp-edged. tins area suffused with pale side. venter entirely smooth except for postanal granules. No inguinal or lumbar gland. Body fairly elongate.. tibia 10. widening on the sides . femur. interorbital diameter equal to that of upper eyelid but only two-thirds the distance between the nostrils. loreal region flat. in postaxillary region a trifle narrower than greatest head width. heels just touch. a small inner and a very small outer metatarsal tubercle. Skin heavily pustular above. 5. Fingers long. toes faintly webbed. with a few weak tubercles in the lumbar region. 7 mm. Color in alcohol. Eye very large and prominent.5 mm.5 mm. third toe much longer than fifth. disk of third finger covering one-fourth the tympanum. with distinct disks. head mm.5 — Head and body.. subarticular tubercles of toes smaller but a little more distinct than those of fingers. a whitish area sharply contrasting with and defining the upper and lower borders of this dark stripe. When hind legs are adpressed. grey. heel reaches anterior corner of eye. indistinct sepia markings between the eyes and down center of back. their distance from end of snout about one-half their distance from eye. a slight ridge behind the tympanum. a large indistinct palmar callus. 9 mm. a short tarsal ridge extending from base of inner metatarsal tubercle and ending in a diagonal gland across the tarsus midway to heel.. passing through loreal regions.. —Dorsum dull heliotrope purple. the upper lip almost in the same plane with it. stripe beginning at tip of snout. at least on its anterior and lower edges. and rather easily torn. scarcely ridged laterally. a sepia lateral and tympanic and continuing to the groin. Tympanum fairly distinct. and vertical. then disks well developed. its greatest width about one-half the diameter of eye and located immediately behind eye. separated from each other by an interval equal to 1% times their distance from eye. 22. Dimensions. first finger slightly longer than second. No external vocal sacs apparent in the male. anterior femur with a dark stripe.. its diameter 1% times its distance from end of snout. head length. hand.FROGS OF COLOMBIA —COCHRAN AND GOIN 49 below the canthal angle. no ventral disk. reaching halfway on antepenultimate phalanx of fourth.5 mm. when limbs are laid along the knee and elbow greatly overlap. a small protuberance of skin above the anus. foot. subarticular tubercles of fingers large but not very sharply defined.

lip white. on lower posterior femur. and runs from the eye to the sacrum. the disks darker below. The skin on back and belly may be strongly or weakly granular. that species is not included in this study. Furthermore. NATIONAL MUSEUM BULLETIN 2 88 posterior femur with a less-defined light dark diagonal crossbars on upper surfaces of femur. and fresh topotypic material that might be considered as representing this species is scarce. Remarks. extending about one eighth the length of the free part set off by a white area below. below knee.50 U. Because of the traces of webs between the toes and of many other similarities. skin tears rather easily. the original description of which states distinctly that the toes are free. and at El Centro. while the maximum size among the 206 examples that were measured for the present study and that here are called brunneus is 28. the full-grown brunneus does not appear to exceed 22.5 millimeters. stripe. although not as easily as that of and P. Dunn (1957) likened brunneus to talamancae. tibia. a sharp dark line along posterior surface of arm. venter white. in Santander. immaculate (in this example) except for a few small spots across the throat. The toes may be faintly webbed at the base. several — of the fourth toe.). chest. hence. but some specimens are entirely smooth below. by the name of brunneus (Cope) instead of latinasus Cope. the size of the type of latinasus was given as three centimeters (30 mm. Sometimes the entire throat. Also. he stated. and foot. it is of some significance when considered along with other facts regarding latinasus. The heel reaches to the center or the posterior corner most examples at hand and it to the anterior corner of reaches to the nostril in one specimen. Considerable variation is noted in the ventral coloring of brunneus as here understood. in Cauca. sharply outlining the dark stripe above it. brunneus each other in the key. a Costa Kican species which. or the webbing may be rather conspicuous. He noted that "in both frogs the white dorso-lateral stripe is entirely above the dark lateral band. has appeared in Colombia at Popayan. of the e}7 e in The the eye in several. The type of latinasus has been missing since Cope's time. with dark irregular spots on upper arm surface." The present authors have not found any specimens of talamancae in Colombia so far.5 millimeters. and belly . While P. Colombia. they are readily separated by the much greater webbing of the toes in palmatus and the relatively shorter femur and hand in brunneus. Although this difference is not great. it seems reasonable to call the very prolific and wideranging little frog found from Panama to Narino. and on foot. Soles of feet and palms of hands pale sepia. palmatus are near Phyllobates palmatus.S.5 millimeters and palmatus grows to a length of 36.

Munchique. chest. In latinasus this light line is said to continue parallel to the dark stripe without crossing — it both becoming diagonal on the side and sometimes extends on the upper proximal part of the femur as a light bar. Occasionally a thin light stripe commences at the middle of the wide dark lateral stripe and continues straight back to the groin. 5. Rio San Juan. 30S0S. 54542-4. its foot is much longer. has much larger black spots all over the lower surfaces.100 ft.7.10. 44044. 1932. CNHM Choc6: Cundinamarca: North 81858-9. ANSP 25353-63.FROGS OF COLOMBIA —COCHRAN AND GOIN are immaculate pale yellow or olive. near Popayan. boulengeri. Selva. 50. Pueblo Rico.860 m. 54496-9. Also. 54339. The white (yellow in life) stripe that starts behind the eye (or sometimes on the canthus) in brunneus begins to curve downward when it reaches the sacrum and goes diagonally across the dark lateral stripe to the anterior surface of the femur. 54545-63. USNM CNHM 150720-1. 54509-26. USNM BM CNHM USNM CNHM CNHM 4336. Medellin. brunneus apparently is larger. 44030-40. 54420-1. Zuaraza. Pueblo Rico. 146466. the entire chin. 2. and its hand is slightly longer. however. of Alban. Villa Maria. CNHM ANSP CNHM 24112-35. near Salento.11. Popayan. Huisito. In femoralis. MLS 139. CNHM 54343. USNM. 146860-2. and sometimes dark dots extend over most or all of the ventral surfaces. Boyaca: Caldas: MCZ 7778-7802.. and sometimes a third light stripe bounds the lower edge of the dark lateral stripe at the lateroventral area in latinasus. . 139b. 20 km. Specimens Examined COLOMBIA Antioquia: Berrio. often enlarging to a definite light spot in the upper part of the groin. Such markings are absent or indefinite in — brunneus. ANSP 25383-7. Caqueta: Aserrio. MLS 162. 44079. 54345. 1910. 54422-7. Pizarro. 51 some fine dark spots occur on the chin. brunneus resembles latinasus considerably in color and general appearance. Laguneta. P. MCZ 2891. CNHM El Tambo. 44001-8. 54543. San Mateo. P. 43997. Quintana. 146867-71.4654491-5. but its head is proportionately a little shorter and narrower. Puerto 63856. Yarumal. 44010-28. Cauca River. 54565-7. irregular black patches on either side of it.. La CNHM 54528-41. while dark spots on a light ground occur only on the posterior half of the belly and on the lower surfaces of the limbs. concentrating on the throat around a light longitudinal median area with large. San Juan slope. 57411. USNM 152320-32. a species of about the same size that might be confused with brunneus. 54457. 5006-7. often. south-southeast of Manizales. 54501-6. and anterior half of belly are solid black. ANSP 25379-82.77-8. the other species with a dark-spotted venter. Cauca: CNHM CNHM Andagoya. Moscopan. 54486. 54429-55.2.

Phyllobates — — — 1944d. Stebbins and Hendrickson.— Dunn. 1957. Hyloxalus palmatus.— Gorham. disk of fourth toe covering about . 146904-20. 1957. free. near Bogota. 54458-64. Prostherapis palmatus. p. 403. Dunn. MLS 410. 1944a. third toe longer than fifth. with lateral ridges. 25.third the tympanum. Dunn. its disk reaching halfway on antepenultimate phalanx of fourth. Toes moderately long. PANAMA: Cans. — — USNM —Stebbins and Hendrick- Description. 401. disk of third finger covering about one.1. USNM 54174. 54342. snout straight-sided. metacarpal tubercles well developed. NariSo: Guitarilla.. Cundinamarca. 150169. CAS 54736. 54466-71. Barbour and Noble. its greatest diameter one-half that of eye.4-7. 150170. 54479. 1959. its lower margin deeply concave before it joins the vertical upper lip. road 150761. interorbital diameter about 1% times that of upper eyelid. 1944. p. Tongue cordiform. 1920. first and second fingers subequal. p. La Guayacana. Can thus rostralis rounded. a little less than interval between nostrils. Magdalena BM Forest. prominent. 1939. 54484-5.1. two-third as wide as mouth opening. 1944b. p. 2. Phyllobates palmatus Werner Plate 9c-e 1899. p. Hellmich. 79.700 m. 1944c. 1957. Cerro Azul. 536. p. its diameter nearly equal to its distance from tip of snout. CNHM 54475-7.2. 146899. 1919. Meta: USNM 81365. Fingers moderately long. a small oval thumb pad present. rounded in profile. 54473. an adult from Buenavista. Buenavista. 118718-9. USNM San Lorenzo. 54185. Meta. 61820. p.300-2. Tuquerres. Santa Marta 1929. (Hypodictyon) palmatus Werner. 61787-8. son. Colombia) Syrrhophus palmatus. Fusagasuga. 1944a. Colombia). Prostherapis granuliventris. Eye large. 77. projecting. 61824-5. Tympanum fairly distinct. 1919b. Bogota. 79. 81 (type locality. Nostrils lateral. separated from eye by an interval equal to one-third its own diameter. Hylixalus granuliventris Boulenger. the upper jaw extending slightly beyond the lower. CNHM CNHM ECUADOR: Quimbo. BM 98. its tip bluntly rounded when viewed from above. 478 (type locality. 54483. 398. 512. loreal region flat. 1939. Hyloxalus granuliventris. Colombia. p. 519. its posterior border free and deeply notched. 77. USNM CNHM de La Macarena. 63005. 1963. to 2 88 San Lorenzo. p.S. p. fig. p. 54487-90. 66318. — — 1959. p. 512. p. 54481. Serrania CNHM 81811-5. fourth finger longer than either and reaching midway on penultimate phalanx of third. p. 52 U.3.2. half-webbed. NATIONAL MUSEUM BULLETIN Magdalena: Finca La Granja. Darien. USNM 54231. 1920. 1957. 1957. Cundinamarca.. Dunn. 1. 77. a larger round palmar callus. oval. Villavicencio. their distance from end of snout equal to one-half their distance from eye. Mountains. p. 54198.

5 mm. no skinfold across the chest. a distinct oval inner metatarsal tubercle when limbs are laid along the sides. from the north to the south of Colombia and from east of the Eastern Cordillera to the valley between the Central and the Western Cordilleras. with some heavy pustules along the sides and on proximal parts of upper femur. ending diagonally across tarsus about two-thirds the distance from metatarsal tubercle to heel. color. with moderately large a narrow wavy ir- sepia interorbital band. as it varies only from 4. Color in alcohol —Dorsum fawn regular sepia spots scattered over it. heel reaches one-third the finger. This character may prove to be of value in suggesting relationships. a heavy skinfold on heel and knee. hand. when hind legs are bent at right angles to body. which probably do not occur in life. On the contrary. few preserved specimens come out of the collector's tin in perfect condition.... a few coarser granules on lower femur. a trace of which appears behind eye and for a short distance along sides. 12 mm. 36. their disks and tubercles pale sepia. with the exception of a wide sepia stripe from tip of snout to anterior eye. When hind leg is adpressed. side of head fawn color. head length. in the males.8 to 5. Dorsal skin so loose that it can be rubbed to resemble longitudinal dorsal folds. as it occurs also in other species somewhat similar to P. head mm. a glandular swelling from posterior corner of eye above tympanum. consequency. Over its wide range. palms of hands and soles of feet ecru drab. Dimensions width.. A pair of lateral vocal sacs and sometimes a widening of the third finger to form a "flange" posterior border of eye. palmatus in structure and appearance.5 mm. throat and chest minutely pustular.FROGS OF COLOMBIA — COCHRAN AND GOIN 53 tympanum and approximately equal to disk of third and a smaller round outer one. tibia. a heavy tarsal ridge. Skin of upper parts delicate and very easily torn.5 mm. no ventral disk. this species is remarkably stable. posterior femur drab with a coarse sepia reticulation. foot. outer surfaces of feet and hands ecru drab. 13. 17. Body very stout. and a transverse irregular spot across snout behind nostrils. the pustules changing to fine granules on posterior part of belly. a considerable variation in the critical measurements would have been expected. upper limb surfaces drab with irregular wood brown cross bars. knee and elbow overlap. 15. heels are separated.5 millimeters in the six measurements that were taken on 105 individuals. femur. no light diagonal stripe on side. — Remarks The dorsal skin is extremely fragile and tears readily. 17 mm. in postaxillary region equal to greatest width of head. ending above shoulder. a pair of lateral external vocal sacs in the male. smooth. 10 — Head and body. anterior femur and entire venter immaculate ecru drab.. with faint sepia spots.5 mm. 337-262—70 5 ..

has no trace of a tail. giving evidence of a pair of lateral vocal sacs. NATIONAL MUSEUM BULLETIN 2 88 The species is quite easy to recognize.5 millimeters still remains. Meta. Some examples have a pale stripe beginning above the shoulder as a narrow light line but widening as it goes diagonally downwards to the front of the groin. in 15 percent it reaches to a point between eye and the tip of the snout. Metamorphosis often takes place when the young are some 15 millimeters in length. Its arms and legs are like those of an adult. Adult males have a dark gray or black throat. This light stripe may have been orange or yellow in life. and sometimes breaking up into several short pale spots. measuring only 12. The torn places in the skin are white. upper Rio San Jorge. and outside of foot. with little of the delicacy of form or pattern found in latinasus. In another 15-millimeter example (USNM 145028).54 U. and some of the other forms in the genus. hence. 150693. Yarumal. and the mouth is not yet that of an 61823 from adult. is 15 millimeters long and has no trace of a tail. The third finger showed a "flange" in only two males (CNHM 61176.5 millimeters. Caldas. Bolivar. and in the remaining 15 percent it reaches to a point between shoulder and posterior corner of eye. and has well-developed limbs and mouth. but often the lighter individuals show the wide darker dorsal chevrons and three wide dark crossbands on femur. . Bolivar. but usually there is at least one small pale spot just in front of the groin. a tail measuring 8. and its mouth is nearly so. MLS USNM 23. often finely or faintly one-third to one-half the so. tibia. from the same place. Some frogs lack the light lateral stripe. The posterior half of the belly is granular. the developmental rate in different individuals may differ considerably. The body color in most preserved adults is a fairly uniform dark purplish brown. 61180) both from Catival. In about 70 percent of the examples measured the adpressed heel reaches to the center or anterior border of eye. UNMS 145030 from Buenavista. for preserved examples are ragged-looking because of the usually torn skin and often are dark colored. is CNHM Specimens Examined COLOMBIA Amazon as: Antioquia: Puerto Narifio. and at a short distance they may give the quite misleading appearance of a lightspotted animal. The toes are webbed in nearly all examples of palmatus. and sometime there a short longitudinal fold down each side of the throat. But Alto de Quimari. brunneus. although the limbs are fully formed.S. and web is conspicuous because of its paler color between the dark toes. USNM 146857-9.

— Breder. USNM 146898-903. 61783-5. Paramo de Verjon.. 1920. p. del Salto Tequendama. 1957. p. MLS 281. 61771.— 1914. 3 (type locality. 61821-3. Finca El Cuchero. 145048-52. p. Quetame. . a short large. Tympanum distinct. ANSP 24322-3. snout very broadly rounded when viewed from above. loreal region vertical. prominent. pp. BM Description. below Tequendama. 643. Honda. Meta: 1329S7. Caqueta: Florencia. furrow between it and the upper lip. distinctly projecting. oval. Tolima: Guandalay. its greatest diameter about half that of eye. an adult female from Pefia Lisa.77. upper 61176-80. [Antioquia. Colombia.. 13472-3. 146874-5. 407. Santa In6s. Narino: La Guayacana. Norte de Santander: La Selva. 150195-8. MLS 153. 1924. their distance from end of snout about one-half their distance from eye. 9-10 (specimens from Santa Cecilia. Camp USNM 145005-15. La Macarena. Las 300 150251. 145032-47. La Macarena. Santander. west of Alb an.21. Monte Redondo. east base of cordillera and extreme limit of llanos. 350-500 m. 78.5.700 m. 145016-31. interorbital diameter 1% times that of upper eyelid. Condoto. Taylor. 1952. separated from eye by an interval slightly concave. Volcanes.S00 ft. a little greater than interval between nostrils. 2. p. 25. 1924 (part). 20425-33. Planta E16ctrica Cundinamarca: Aguadita. Dunn. . on road between Sibate and Aguadita. 391. just off edge of Bogota savannah. MLS 136. 11. 399. Rio San Jorge. 150199-250. Santander: Mogotes. the upper jaw extending well beyond the lower. Costa Rica. BolIvar: Alto de Quimari.— Dunn. 146840-2 (juv. below Tequendama Falls. Santandercito. east of Bogota. 1. Velez. north of Medellin. 1946.) 1913. p. 274. Sabana de Bogota.500 m.] Colombia. 146893-7. USNM 146937-8. USNM m. 145053-4. Choco.000 CNHM CNHM USNM AMNH USNM USNM AMNH AMNH USNM AMNH USNM AMNH USNM USNM USNM USNM USNM USNM USNM USNM CNHM m. Tongue half as wide as mouth opening. 150187-93.— Barbour and Noble. 137773-86. 1963. 5-7. its diameter equal to its distance from tip of snout... 20409-18.— Gorham. p. 22610-5. Bucnavista. CNHM 61776. 20436-7. Foineque. USNM Phyllobates pratti Boulenger Plate 11g-i 1899. 145055-7. Eye prominent. 20372-3. rounded in profile. 118720. near Tocaima. Quebrada La Lechera. 144998-5004. Bogota. 3. Arracachal. — 389. Catival. (This locality record has been questioned by Dunn).— COCHRAN FROGS OF COLOMBIA AND GOIN 55 USNM Atlantico: Barranquilla.200-2..). and Panama). AMNH 20359-63. 1. pp. MLS 224.— p. 2. San Gil. its posterior border free and unnotched. Choachf. Canthus rostralis straight. Phyllobates latinasus (not of Cope). Nostrils lateral. Caparrapi. 118228. 1026. USNM 147132-4. 1931. 146634. Phyllobates pratti Boulenger. 150194. 281a. fig. above Camp Ranas. p. Tequendama Falls. 20364-9. 145059-71.

metacarpal tubercles well developed. foot. above tympanum. Most examples of Phyllobates that could be related to P. in postaxillary region less than greatest width of head. a white (yellow commencing behind eye and extending straight back to the upper part of the groin. have this stripe going straight back from above the shoulder so that it ends above the hind leg insertion. head mm. 7 mm.. but a few frogs. nearly immaculate. heels are narrowly separated. a weak ventral disk. a heavy skinfold on heel and knee.5 — Dorsum immaculate Mars brown. Dimensions. including chin and chest.. disk of third reaching to base of antepenidtimate phalanx of fourth. pratti . smooth (faintly pustular under the lens) venter smooth a distinct glandular dorsolateral ridge from posterior corner of eye.56 U. second and fourth subequal and reaching base of penultimate phalanx of third.5 mm. soles and palms drab with indistinct darker marblings. immaculate pale olive-buff. 5 — Head and body. knee and elbow touch.5 mm. continuing on loreal region. a light stripe along upper venter. 6. a pair of lateral external vocal sacs in the male. with lateral and small terminal disks. 6.. Fingers long. slightly (about one-eighth) webbed. Toes fairly long. an oval thumb pad present. where it merges with the light buff upper surface of the femur. Color in alcohol. posterior femur buff. as further lip. — observations on the significance of direction in the light lateral stripe are needed.. NATIONAL MUSEUM BULLETIN 288 equal to one-fourth its own diameter. 18 mm. pratti have this stripe dropping diagonally across the side to the lower part of the groin. with a large oval sepia spot from its midpoint to behind the knee. when hind legs are bent at right angles to body. third toe much longer than fifth. 8. both following the light longitudinal ridges . heel reaches center of eye. Remarks. stripes. and with a few lighter spots behind anus and extending along lower femur to knee. tibia.S. head length. width. and their disks small.5 mm. when limbs are laid along the sides. Skin of upper parts rather easily torn. a skinfold across the chest. resembling the examples of from the British Museum. femur. a small tubercle on tarsus midway to heel. hand. and forearm pale wood brown with darker bars and spots. Body elongate. . upper part of femur pale. first finger much longer than second and equal to third. widening in life) dorsolateral longitudinal line behind the eye and along the side to groin. a small round metatarsal tubercle and an outer one of equal size. ending at groin. The validity of this species is still doubtful. foot. and another such ridge below it. with a wavy but sharp demarcation between it and the clove brown lateral band. free.. upper part of tibia. a clove brown band beginning at tip of snout.. mm. no apparent tarsal ridge. 7. a large rounded palmar callus. When hind leg is adpressed.

west of Playa de Oro. p.45.10. Colostethus latinasus Cope. a small oval inner metatarsal tubercle and a smaller round outer one. sharp tarsal ridge. ending diagonally to heel. reaching to base of penultimate phalanx of third. 1924. 130. prominent. but a comparison of pratti should Boulenger's material of the former with freshly collected specimens of the latter shows that there is slight difference between the two. when limbs are laid along the sides. slightly projecting. p. 10 km. rounded in profile. Barbour and Noble (part).30. 1920. p. the disk of which covers about one-fourth the tympanum. p. — Granada). an adult from Montanita. 1963. Juan. fourth longer than either. — Cope. Body rather elongate. p. 25. 1910. Rio San Juan. distinctly webbed at base. Specimens Examined COLOMBIA Noanama.12. knee and elbow overlap. with lateral ridges.39. heel reaches center of eye. first finger a little longer than second. USNM 150716. it seems advisable to retain Boulenger's name. 399.80.— Gorham. loreal region flat. interorbital diameter V/2 times that of upper eyelid. Rfo Tamana of Rfo San BM 1913. rostralis upper lip. a distinct skinfold hind leg is adpressed. p. — Description.5. its greatest diameter about half that of eye. snout very broadly obtuse when viewed from above. 1957. 119. Colombia. 131. Pefia Lisa. p. When short. a larger round palmar callus. pp. 1866. 78. Colostethus latinosus. — New Truando River. Condoto.11. a halfway on heel and knee. Tympanum distinct. 1898a. flaring outwards to the Eye large. Choc6: BM BM 1909. 1866. — Boulenger. Caldas. USNM 147208-9.FROGS OF COLOMBIA —COCHRAN AND GOIN 57 Barbour (Barbour and Noble. 399) persuaded Boulenger that be synonymized with latinasus. Canthus rounded. 1874.21. p. 1874b. 1954. Novita. narrower than interval between nostrils. their distance from end of snout one-fourth their distance from eye. metacarpal tubercles well developed. 1914.— Dunn. its posterior border free and unnotched. Boulenger. For the present. 119. a small oval thumb pad present. PJiyllohates latinasus Cope Plate 10g-i Phyllobates latinasus Cope. third toe longer than fifth. Tongue nearly two-thirds as wide as mouth opening. when hind legs are bent . Toes long.. 1898a.— Griffiths.77. 1920. the upper jaw extending well beyond the lower. Nostrils lateral. its disk reaching to base of penultimate phalanx of fourth. 130. oval.11. 41. its diameter about four-fifths its distance from tip of snout. faintly webbed at the base. 1863. separated from eye by an interval equal to one-fourth its own diameter. 1866b. Fingers long and slender. 300 ft.7. p. in postaxillary region narrower than greatest width of head. 2. 5. disk of fourth toe covering one-fourth the tympanum and equal to that of third finger. 48 (type locality. p. vertical. p.

. Barbour and Noble (1920. widening behind eye. irregular white line goes forward along the side from the feet slate-gray. although a graph of critical measurements indicates a slight overlapping with that of mertensi. apparently no external vocal sacs in the male. .5 — Head and body. straight parallel lateral line have come to light. immaculate except for a black line around lower jaw and a patch of dark dots below shoulder. pi. 1899. 9 mm. and Narino. with an irregular and tubercles a of hands and A narrow. no ventral disk. 11 mm. femur. heels touch. 11. these are from Choc6.S. below by sepia spots. 8 mm.- width. 10. 399) believed latinasus to be conspecific with Boulenger's pratti. p. venter stripe edged white on a pale buff. 274. The foot of latinasus averages proportionately shorter than that of any other Colombian species of Phyllobates. 26 mm. and Dunn as to what latinasus really might be. it is very difficult to identify any frog positively as Phyllobates latinasus. Four examples that do have the non-oblique dorsolateral line and the short. groin. foot.58 U. 3). Even the example chosen for the preceding redescription does not agree entirely with the earlier conceptions of Barbour. — Dorsum clove brown.5 mm. and the present lack of freshly collected material from the type locality. the disks little darker. p. NATIONAL MUSEUM BULLETIN 288 at right angles to body. row of black spots preceding posterior femur.. in which the light dorsolateral stripe continues straight back to the groin. continuing on loreal region.. hand. lower surfaces anterior femur sepia. and above it a semilunar pale spot from the groin halfway to knee. upper lip cream buff below the black lateral stripe. Boulenger. Although Cope states that his example had an oblique white line from the femur along the side.. outlined above by a seal brown dorsolateral stripe which lightens at midbody to white (yellow in life) anterior femur with a clove brown stripe. remainder of Color in alcohol. 12 mm. and dropping obliquely to groin. pustular anteriorly. tibia. a black lateral stripe beginning at tip of snout. with a short parallel white stripe along the side from groin forwards nearly to axilla. Colombia. the faded and softened condition of the type specimen from Truando at the time of description. head mm.. Skin of upper parts easily abraded. a swelling behind posterior corner of eye above tympanum. with large conical tubercles on sacrum and upper surfaces of legs. a weak skinfold across the chest. Remarks. 7. head length. below the dorsolateral stripe. Valle. —With Cope's wholly inadequate description. Dimensions. . powdered with small sepia dots.. venter smooth (minutely pustular under the lens). fig. as shown in the figure of pratti (Boulenger.

Goin. near Cana. Hyloxalus fuliginosus Jiminez de la Espada. FROGS OF COLOMBIA The upper femoral — COCHRAN AND GOIN 59 surface of latinasus tends to be pale. 54486. p. 54542-4. ft. and very all.1. 1935. 8.4-7. Pueblo Rico.7. Montafiita. 81858-9. west of Playa de Oro. El Tambo. 1871.. Quintana. ProsLherapis fuliginosus.000 3. 5. Novita.— Breder. Medellin. 66318. 1935. ANSP 21797-800. 44044. 1963. Cauca: CNHM CNHM CNHM CNHM CNHM BM CNHM BM BM CNHM USNM USNM USNM USNM USNM (tentatively identified) Valle: BM BM 95.6.200ft. 147430. — — 77.12. a slightly mottled. south-southeast of Manizales. Cauca River. 54550-3. BM 98. west of El Tambo.. p. BM 1932.7.— Gorham.29]. 54174. lower Calima River. 1875. 1914. 155. la-b (no text) .77-8.3.10. but on the posterior surface is a sharply marked.] MLS 35. Rio Rosario. 437. 54565-7. CNHM 43997. Pasto. 59 (type locality. nine km. 24259-60. Rio Grande. Pefia Lisa. 92595. USNM ANSP USNM 54231. Pueblo Rico. 54555-63.11. [Tentatively identified: Montafiita.21. EQUADOR: PANAMA: Quimbo. ANSP 25353-63. USNM Near Buenaventura. 167. 44001-8. — — Evans. La Selva. and also ends at the knee. p.11. 1942. San Jos6 de Moti.11.80.—Myers. Ecuador). ANSP 25379-82. p. CNHM 54496-9. 147487. 54470-1. MCZ 2891. 1946. MCZ 16083-8. 150 ft. 20 km. Breder.. 405. 1960. 147208-9. NariSo: Imbili. 54509-11. Dunn. Yarumal. Rio Mira. Moscopan. crosses the upper proximal part of the femur. USNM Cerro Azul. 162.. 54339. — .11. Munchique. Zuaraza.. p.10. 5. east of Santa Rosa. San Juan. 95. 10 to 15 km.46-50. Pacora. 1909. 54429-55. 405. USNM 150715. 54345.39. 1960b. 3893. figs. Colombia]). USNM 146466. Cana. Popayan. 6006-7. Rio San Juan. 1947. if at close resemblance. Caldas: Pueblo Rico.2. Guisito [Huisito]. 54545-6. and below that stripe is a wide white stripe that begins at the groin. 63005. 54457. 1913.11. p. 1910.77. 44079.30. 137 (type locality. 146248. pi. Rio San Juan. 54422-7. 25. west of Pasto. Pizarro. 506. 54501-6. USNM 145793 [formerly Carton de Colombia. 1957. BM 1910. p.16. p. Prosthcrapis inguinalis Cope. [Choc6]. 54543. ANSP 25383-7. This light stripe is bordered below by a very irregular dark stripe that breaks into small and large spots on the lower inferior part of the femur. 44010-40.860 m. MLS 391. ft. Rio San Juan. Condoto. Phijllobates inguinalis. USNM 150716-7. Specimens Examined COLOMBIA Antioquia: Sonson. El Valle.45-6.100 ft. CNHM 54491-5. Noanama. A comparison of this species with brunneus shows in general. wide black stripe ending at the knee. USNM 150718-9. Parker. Truando River. Gatun. USNM Phyllobates inguinalis (Cope) Plate 10a-f 1868. p. 2.5. Rio Tamana. Laguneta. Choc6: Andagoya. 146469 North of Tumaco. 44047. New Granada. Camp 149722. MCZ 7778-802.. 3. 1946. p. 54336.30-1.. ANSP 24112- CNHM 63856.300 Cauca River side.

followed by a narrow vertical cinnamon bar. Colombia. a round palmar callus. separated from eye by an interval equal to one-third its own diameter. axilla white. with small drab spots near border of lip. the upper jaw extending well beyond the lower. a swollen area above no ventral disk. but with edge of lip unspotted ecru. Eye large. bluntly rounded at the tip when viewed from above. from Truando River. a wide dark part of its original pattern can be discerned. venter smooth. 28.. its posterior border free and unnotched snout straight-sided. a few pustules on snout. ending way and marked by an elongate tubercle at that point. 12.5 mm. and along side of body. equal to interval between nostrils. first finger a trifle longer than second and fourth and reaching to base of penultimate phalanx of third. heel reaches center of eye.S. a pair of lateral external vocal sacs and sometimes "flanges" on the third finger.5 mm... — hand. 7 mm. its disk reaching to center of antepenultimate phalanx of fourth. interorbital diameter slightly greater than that of upper . oval. head width. 2 88 — USNM Description. loreal region concave. 4349 (holotype).5 mm. tibia. 9. a head and anterior half of back pale russet to wood brown. Head and body. Fingers rather short. femur. 9 mm. a slight tarsal ridge. oval inner metatarsal tubercle and a smaller diagonal outer one. the chest. a heavy Body moderately stout. Nostrils lateral. with traces of webs at the base and with strong lateral ridges. angles to body. behind ear. metacarpal tubercles well developed. scarcely projecting. When hind one-third the to heel skinfold on heel and knee. in postaxillary region narrower than greatest width of head.60 NATIONAL MUSEUM BULLETIN U. knee and elbow touch. upper lip russet. Choc6. their distance from end of snout one-fourth their distance from eye. Skin of upper parts easily torn. its diameter equal to its distance from tip of snout. 12. sacral region ecru drab with irregular traces of russet in the middle. a small. in the male. nearly smooth. prominent. Dimensions. an indistinct oval thumb pad present. Color in alcohol. Toes moderately long. the sides. one-fourth webbed. its greatest diameter apparently about two-fifths that of eye. with some glandules on lower and posterior tympanum. —Although the type specimen Top is much faded. disk of fourth toe covering about one-half the typanum and noticeably larger than that of third finger. 11 mm. third toe a little longer than fifth. heels are separated.. Tongue half as wide as mouth opening.5 mm. head length. furrowed at its junction with the nearly vertical upper lip. truncate and sloping slightly backward in profile.. the disk of which covers about one-fourth the tympanum.. of . eyelid. foot. Tympanum indistinct. no skinfold across surfaces of femur. when limbs are laid along when hind legs are bent at right leg is adpressed. Can thus rostralis rounded.

The white stripe often continues along the front of the femur. 150174. 150176-7) have the than the second. sometimes narrowing and extending to the shoulder. and sometimes it is heavily spotted over the entire belly. The light lateral stripe in front of the groin is most noticeable. the second between the shoulders. although the limbs are fully formed. While some freshly preserved frogs are more or less clove brown. posterior femur tibia. with some large dark spots on the lower outer limb surfaces. disks. and forearm covered by coarse dark reticulations extending to upper limb surfaces as dark crossbands.5 millimeters long. — — Remarks. its line. soles of feet and palms of hands russet. showing three darker chevrons on the back the first chevron between the upper eyelids. The venter is smooth in nearly all examples at hand. This light extension lies across the center of a wide dark longitudinal stripe originating on the tip of the snout and continuing to the groin." The webbing of the toes is quite variable being a mere trace in some frogs but extending for about one-fourth the length of the fourth toe in others. The adpressed heel reaches the center of the eye in half the frogs examined and other half. a pale ecru stripe continued on anterior proximal surface of femur. apparently lateral band. although a few show granules on the lower femur and beside the vent. A young specimen from El Centro (USNM 150185) that measures 10 millimeters in head and body length is fully metamorphosed. sometimes breaking into a series of small light dots. An 8-millimeter tadpole from the same place (USNM 150186) has a tail 9. with a few cinnamon spots on lower femur and tibia. to the anterior or the posterior corner of the eye in the The first finger is longer considerably and sometimes slightly (USNM so.—COCHRAN FROGS OF COLOMBIA AND GOIN 61 sinuous lower margin emphasized by a narrow darker beginning behind eye and bordering the dark lateral stripe above. and webs paler. others are light brown. . Sometimes the venter is nearly immaculate except for some scat- tered dark spots around the chin. venter ecru drab. having no tail or other larval characteristics. although it may be interrupted by the dark femoral crossbands. then descending diagonally across it and continuing as a pale elongate spot just in front of groin. On lighter examples this dark stripe often is apparent. and the third on the sacrum. but sometimes it is suggested only by small light and dark spots along the side on both borders of the lateral stripe. sometimes Several males from El Centro membranes of the third finger widened into large "flanges. Nearly all preserved specimens show abrasions of the skin. the tubercles.

USNM 139710. 1. 1963. Azul. 73. with a slight furrow between it and the upper lip. Dunn. 1944a. USNM 50227. slightly projecting. 2. 150719.150 ft.000 ft. Boquer6n of Bogota. 401. 81752-8. near Valle de Ant6n.— Gorham. 81887.000-2.. El Valle. the upper jaw extending well beyond the lower. region. 1957.S30- La Uni6n. Chingasa. 146248. 1914.] Colombia). all in Cundinamarca. 144995. 503.] Colombia). 1926b.. with bulbous tips. an adult female from Bogota. 1920. Barbour and Noble. 1. km. Tongue slightly more than half as wide as mouth opening. 2. 73. ANSP 22557. 1924. Prostherapis subpunctatus Cope. p. fig. 150171-86. 150692. p. Caldas: USNM USNM USNM USNM USNM USNM USNM CNHM USNM Santa Elena. 1920. interorbital diameter 1% times that of upper eyelid. 140659-60. Cana. 2.. 73.300 m. 1944c. Colombia. Eye large. oval. p. Jacinto. of — 1944a. Prostherapis tarsalis Werner. east of Santa Rosa de Cabal. 2.. 7. Canthus rostralis distinct. USNM 129901-4. Prostherapis variabilis Werner. Bogota. USNM 139900-4. CundiDescription.860 m. 146929-36. ANSP 22494.050 ft.800 m. 2.— Noble. 2. Peracca.. 9. USNM 146466. probably upper Sinii River. 1959. 147125-31. Choc6: La Granja. p. separated from eye by an interval equal to one-third its own diameter. 103. Veragua.— Stebbins and fig. [Cundinamarca. 1939.. ANSP 21738-9. Tres Puntas. p. Dunn. 1. 473 (type localities. Bogotd. Truando. prominent. Sarare Norte de Santander: La Selva. 8 (reprint) . 5. equal to interval between nostrils. 25. Paramo of Fusagasuga). Herrera. Bogota [Cundinamarca. Nostrils lateral. 1944a. road to Choachi. 139691-3. Fusagasugd. 1899. p. Colombia: Alto von Sibate. NATIONAL MUSEUM BULLETIN 288 Specimens Examined COLOMBIA BolIvar: Alto de Quimarf. Dunn. Tympanum very distinct. Cerro ANSP 22543-4. La Loma.. nine 20 km. USNM 146942-51. 520. Narino: West of Pasto.—Dunn. 1. 146939-41. El Valle de Ant6n. La Campana. Pasto.800 ft. p. Rio Cana. its diameter equal to its distance from tip of snout. p. its greatest diameter two-thirds that of eye. p. 2 (type locality. Herrera.. Hellmich. south-southeast of Manizales. 303 (type locality. 146469. Santa Librada. Santander: El Centro. Tierra Nigra. — — p.250 ft.62 U. Hendrickson. pi. ANSP 22345-6. snout rounded when viewed from above. 4349 (holotype). 140060. Las Minas. CNHM 61778-82. head of Madden Lake. its posterior border free and very faintly notched. their distance from end of snout nearly equal to their distance from eye. p. USNM CNHM 61827-9. 2. — — — p. Valle: USNM USNM USNM USNM Phyllobates subpunctatus subpunctatus (Cope) Plate 11a-c 1899. Fingers long. truncate and sloping backwards in profile. loreal region flat. Paramo 1916. 118229.S. Boyaca: Leiva. — CNHM 78. 2.. Pequeni River. p.400 m. p. Phyllobates subpunctatus. PANAMA: Cerro Manguillo. . 66319-20. p. namarca.

palms of hands and soles of feet vinaecous-buff. apparently no external vocal sacs. a slight ventral disk. with wide diagonal sepia crossbands. Although Dunn (1944c) wrote that subpunctatus was limited to the general vicinity of BogotA in southeastern Cundinamarca. metacarpal tubercles well developed. with some large seal brown spots on outer parts of femur. its pressed. stripes along the back. venter coarsely granular on belly and lower femur.5 mm.5 mm. a low swelling in parotoid region. the snout pale wood brown in front of this spot. Body moderately slender. the minute webs between the toes nearly white. head length. venter pinkish-buff to vinaceous-buff. 7. 10.5 mm. in postaxillary region narrower than greatest width of head. ridged. a heavy tarsal ridge. a large. minutely granular on sacrum. Skin of upper parts resistant to abrasion. when hind legs are bent at right angles to body. 7. with small scattered sepia spots beginning on anterior belly and ending at level of groin. upper surfaces of limbs wood brown. a wide light dorsolateral stripe from posterior corner of eye diagonally to groin. tibia. free. heel reaches posterior border of tympanum.5 mm. head width. the tip of which covers about one-eighth the tympanum. which reaches to base of penultimate phalanx of third. first finger a little shorter than second. no glandular ridge from posterior corner of eye. he nevertheless identified as this species two examples that he collaid along the sides. the tubercles and disks lighter. upper lip wood brown. and foot. When hind leg is adfifth.FROGS OF COLOMBIA — COCHRAN AND GOIN 63 with lateral ridges. 22 mm. a skinfold on heel and knee.. Toes webbed at the base. third toe much longer than disk reaching midway on antepenultimate phalanx of fourth. and larger than that of third finger. 9. — — — ..5 mm. with darker areas on outer phalanges of foot. Dimensions... lower limb surfaces vinaceous pink. tibia. foot. disk of fourth toe covering nearly one-fourth the tympanum.. a round palmar callus. and sides. posterior femur with the suggestion of a light longitudinal stripe bounded below by a broken and irregular sepia stripe and above by the ends of the diagonal sepia crossbands. when limbs are knee and elbow slightly overlap. Remarks. a distinct oval inner metatarsal tubercle and a smaller round outer one. nearly smooth anteriorly (pustular under the lens). weak thumb pad present. hand. immaculate. 8. bordered below by a slightly narrower sepia stripe whose anterior part commences on the loreal region. 6 mm. long. Dorsum cinnamon. square sepia spot between and behind eyes.. a small. no skinfold across the chest. the inner pair broken up into elongate spots. Head and body. becoming smoother on chest. with four indistinct sepia Color in alcohol. chin. ending diagonally midway across the tarsus in a heavy tubercular ridge. heels fail to touch. femur.

always is present on the femur may be pale to medium brown. p.4 and 7. and the toes webbed at the base (although sometimes very slightly) in the 48 examples at hand from Cundinamarca and Meta. The toes are unwebbed in 18 adults. The measurements falls between 5. and to the anterior corner of the eye coefficient of variation in all critical only one case) in 2 percent. 194) as Liuperus elegans Peters has Museum (BM 68.9 in these examples. Its color is . and it slants downward from above the level of the elbow. and with webs almost one-eighth the length of the fourth toe in three. immaculate or with a few small dark dots. The light dorsolateral stripe beginning behind the eye is wide and distinct on nearly all examples. The first and second fingers are subequal in 24.3.7). while the in the second finger seems to be slightly longer in 18. The heel reaches between the shoulder and the posterior corner of the eye in 74 percent of the frogs. venter may be immaculate pale buff or light olive. or a series of irregular dark markings. indistinguishable from the A Bogota frogs. The heel extends no further than the tympanum in eight and to the center or to the posterior or anterior corner of eye in 33. the back practically smooth. NATIONAL MUSEUM BULLETIN 288 Meta. but sometimes a rather concentrated dark spotting appears on the posterior half of the belly and on the under surface of the femur. or many heavy or medium spots. during his visit of 1943-1944.5 millimeters in length.S. Hno. which is less in the Cauca frogs. The belly is more or less granular. Cauca. to the center of the eye in 24 percent.4. The largest example frogs 151127-79) Cauca series is 23. suggest subpunctatus in many characteristics but vary slightly from Meta and Cundinamarca examples. Niceforo Maria has sent eight other examples from the same locality.64 U. The skin is not abraded in any specimen in the series. Some (USNM from Purace. Their critical measurements agree except for foot length. series of spots nearly Above this the top of the regular border along the sides of the body. mostly near the lateroventral area and along the midline of the lower femur. The sides are dark. all of them seemingly lected in Villavicencio. and these resemble Dunn's specimens closely. slightly webbed in 20. The dark area bordering it above is narrow and often broken up into spots anteriorly. Some examples have a few large conical tubercles above the knee and on the upper surface of tibia and foot. (in The An example from Bogota mentioned by Boulenger as having been identified by Giinther been loaned to us by the British (1882a. ending in the groin. this area being quite wide and ending in an ir- dark stripe or anterior femur. The posterior femur may have a continuous dark stripe like that on the anterior surface. while again only a few dark spots may show.

Choachf. Boyaca.000 m. paramo above Bogota. 142800-4.. 132989-90.1 percent in 13 examples first and second fingers usually subequal. east of Bogota. MCZ 7309-14. Abayo. 146213. or first finger shorter than second. ANSP 24396-7. Arcabuco. Tequendama. west of Bogota. 150711-4. Paramo de Palacio. 2. north of Alban. 1960. 9036-40. USNM 150710. dorsal skin smooth. but with the very short between 37 and 46 percent of total length. MLS 402. Paratypes. 132991-3. Parque Nacional. resistant to tearing or abrading. Monte Redondo. new subspecies Plate 11d-f Holotype. Colombia. 146921-9. 150313-4. 18 km. same data as holotype. 146879-87. 2.000 feet. 3. 145699-705. near Ramiriqui-Rondon Road. belly with round brown spots becoming larger and closer together posteriorly. 68. 14 km. La Mesa.800-2. northeast of Nazaret. Cerro de Torquita. USNM 150701-8. Sabana de Bogota.SUNHM 20290.FROGS OF COLOMBIA —COCHRAN AND GOIN 65 almost gone but otherwise it is in good condition and its measurements and other characteristics prove it to be a true subpunctatus. 145805 (formerly 20303). Peters' true Liuperus elegans definitely has been identified as an Eleutherodactylus and is treated in the present paper as Eleutherodactylus elegans (Peters). El Dintel. Wales. H. 94967-77. collected from under rocks along northeast shore of the lake. Boyaca. 3. 150267. up from Bogota. near Miraflores. Cundinamarca: USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM CNHM CNHM USNM Phyllobates subpunctatus walesi. 150722.3. Boyaca. Meta: Villavicencio. USNM 151874. 150266. USNM 150709. San Miguel. Municipio Chia. — Resembling Phyllobates subpunctatus. 138758. MLS 386. 118715-6. and a Diagnosis. 150295-312. Zetaquia. lower anterior edge of femur with a wide light stripe bordered above and below by dark stripes. Paramo de at about 10. MCZ 9734-9.4.500 m. USNM 150694-700. 145104.. MCZ 16268. from Laguna de Tota.000 m. Peligro. near Bogota. vicinity of Monserrate. 81884-5. Santander. 150265.. tongue not indented. 150262-4 (150264 consists of three tadpoles). east of Bogotd. by J. Tunja. 40 mi. toes nearly one-fourth webbed to less than one-eighth webbed. February 3.800 m. near Bogota. 150252-61. paramo just east of Bogota. Specimens Examined COLOMBIA USNM CNHM BM USNM 146876-8. USNM SUNHM la Rusia. 146888-92. adpressed heel reaching to posterior corner of eye or posterior border of tympanum. east base of Cordillera of Bogotd and extreme limit of llanos. throat and chest immaculate cream buff. 81886-9. averaging 41. below Tequendama. Boquer6n.M 20291-302. —SJJNB. Bogota. Paramo Cruz Verde.. three km. 144996-7. Boyaca. 3.. 150268-94. 118237-8. Boyaca.7.900 m. El Verj6n. tibia — — .

knee and elbow touch. a larger round palmar callus. heel reaches to posterior border of tympanum. a small distinct oval thumb pad. diameter four-fifths its distance from tip of snout. rather prominent outer one: a strong tarsal ridge. external vocal sacs in male. no apparent prominent. with small lateral ridges and faint traces of a web. throat and chest straw yellow with a few sepia spots below chin and a narrow sepia line around lower jaw. nearly one-fourth webbed. a curved black bar between the eyes. loreal region concave. broadly ovate. reaching to center of antepenultimate phalanx of fourth. slightly projecting. Toes moderate. ending in a diagonal swelling midway to heel. dark-bordered stripe on front of forearm. a small but distinct inner metatarsal tubercle and a somewhat larger. a pronounced skinfold on heel and knee. its greatest diameter one-half that of eye. foot. clove brown. Dimensions.S. metacarpal tubercles well developed. 7 mm. in postaxillary region a little wider than greatest width of head. Skin of upper parts not subject to abrasion. continuing below canthus to eye. . — Description of holotype. narrowing to a ridge along side. tibia. granular above anus. When hind leg is adpressed. Fingers moderate in length. 9 mm. some very fine diagonal glandular ridges on side. whose disk covers about one-eighth the tympanum. Nostrils lateral. its posterior border free and unnotched. fourth finger much longer than either and reaching to base of penultimate phalanx of third. hand. head length. where it curves downward in front of femur.5 —Head and body.. the upper lip flaring slightly beneath it. width. including head and upper limb surfaces. bending downwards to end in a semicircular gland just in front of groin. Body rather stout. head femur. when limbs are laid along the sides. size maximum known. 6 mm. 23 millimeters. smooth anteriorly. the disk of which covers about one-fourth the tympanum and is larger than that of third finger. a light stripe from tip. the upper jaw extending well beyond the lower. 10 mm. a glandular swelling from above tympanum. third toe a little longer than fifth. their distance from end of snout two-thirds their distance from eye. —Dorsum.. . its . and following below the narrow glandular line to groin.66 U. when hind legs are bent at right angles to body. . truncate and sloping backwards in profile. mm. 7. Color in alcohol. snout very obtusely pointed when viewed from above. interorbital diameter twice that of upper eyelid and equal to interval between nostrils. venter smooth some weak granules on lower surface of femur. heels touch. . a wide clove brown snout . 23 mm. no skinfold across the chest. Tympanum small but distinct. 9 mm. Eye large. first and second fingers subequal. Can thus rostralis rounded but definite. Tongue as wide as mouth-opening. no ventral disk. NATIONAL MUSEUM BULLETIN 2 88 similar light. separated from eye by an interval equal to onehalf its own diameter.

the remaining half being wood brown with many round dark spots scattered over it. 20298 is paler than the others and has three dark longitudinal lines on the back. thighs and feet of specimens in the series 150711-4 from Tunja. Distinct traces of Saturn red can be seen on the lower surfaces of arms. In all of the Tunja frogs the chin has few to many brown dots. subpunctatus.2 percent. in the others large dark dots appear. in spots on the belly. a larger Remarks. so obvious in most Cundinamara examples. being from 4. The nominate form of subpunctatus is known at present only from Cundinamarca and Meta.3 millimeters. can be made out only one or two specimens of P. these spots being black on the dark area and sepia on the lighter area. and to the The webs on the tympanum or shoulder in 37 percent toes are much less strongly developed in the latter. of P. In one of these. Its tibia is usually longer than that of the Boyaca form. from about one-eighth to nearly one-fourth the length of the fourth toe. The toes usually show distinct traces of webs. are s. then narrowing above axilla and continuing diagonally to groin.3 to 8. however. Average length of tibia in the Boyaca form is 41. and the wide light dorso- lateral stripes. or ventral pattern SUNHM USNM nearly so. is nearly the same in all examples from Laguna de Tota. The Boyacd frogs are darker in color. variation in the critical not excessive. measurements and much The the is different frog. but the throat and chest are immaculate. This latter stripe is not continuous with the one on the lower anterior femur. The heel consequently reaches the eye in 63 percent of true P. . hands. The tibia is proportionately shorter in this subspecies than in any other known Colombian Phyllobates except mertensi. The heel reaches to the posterior border of the tympanum in most specimens and to the posterior corner of the eye in one example. The dark ventral spots are more densely grouped in some.FROGS OF COLOMBIA band across the anterior — COCHRAN AND GOIN 67 half of the belly. the median one encircling a pale diamond-shaped spot behind the occiput and the two outer ones bordered below by a pale stripe beginning in a wide light area covering the glandular swelling behind the eye.7 percent of the total length in the 48 examples measured. averaging 43. s. and the first finger is decidedly shorter than the second in three others and is not fully developed in a juvenile. as a narrow black ring encircles the leg at its insertion. The first and second fingers are subequal in 14 frogs. —In examples at hand. the central part of the belly is immaculate. with a sprinkling of very fine dots around them. and more widely scattered and less evident in a few. walesi. walesi. much more apparent The dark The in walesi.

truncate in profile. — — Gorham. Colombia). Maxillary teeth present but rather weakly developed. ovate. p. reaching to base of penultimate phalanx of third.). a small oval thumb pad present.800 m. distinctly projecting. free. Proslherapis vergeli. slightly notched in the center. All other examples have an immaculate throat or a pale throat with scattered small Tunja show a dark either black spots. its disk reaching to center of ante- . 1. 1940b. 25. Colombia. equal to interval between nostrils. Toes moderately long. Cundinamarca. 963. honor of its collector. which covers about one-third the tympanum. the upper jaw extending well beyond the lower. Tympanum rather indistinct posteriorly. descending vertically to the upper lip. its diameter nearly as great as its distance from tip of snout. a larger rounded palmar callus. Wales. The eight examples from Peligro on the Ramiriqui-Rondon road have relatively longer legs than the other specimens. Can thus rostralis rounded. loreal region nearly flat. its posterior border free. Eye large. first finger shorter than second. with lateral ridges. third toe much longer than fifth. H. near Fusagasuga. in the other examples insertion to the center of the which it it extends from the anterior shoulder tympanum. The collection and study of more extensive material may them serve to differentiate This form is named in positively. prominent. J. NATIONAL MUSEUM BULLETIN 2 88 known maximum size of walesi (25 mm. interorbital diameter IK times that of upper eyelid. as the adpressed heel reaches to the center or to the posterior or anterior corner of the eye. 122. Hyloxalus vergeli Hellmich.5 mm. MLS 309. 1963. a young adult from two kilometers southDescription. although five have black throats (probably a male characteristic) while the other three have light throats that are more or less thickly black-spotted. except in two individuals in reaches the posterior corner of the eye. west of Fusagasuga. Two of the five specimens from suffusion on the chin. Finca El Vergel.third its own diameter.) appears to be a little less than that of the nominate form (27. Nostrils lateral. p. one-half webbed. 2 (type locality. figs. its greatest diameter about one-third that of eye. snout broadly obtuse when viewed from above. Phyllobates vergeli (Hellmich) Plate 9f-h 1940. 1. their distance from end of snout about one-third their distance from eye.S. separated from eye by an interval equal to one.68 U. It is not feasible to give a separate name to the Peligro frogs at this time.. lobate. Fingers moderate in length. Cundinamarca. fourth longer than either. These eight specimens are rather similar in color. metacarpal tubercles well developed. tongue half as wide as mouth opening.

9. The young specimen is darker. 11. 22. as well as some dark spots on the top of the head. smooth to pustular.. ending diagonally midway to heel. when limbs are knee and elbow greatly overlap. but the chevron mark above the shoulders that appears in the figure of the type. side of body pale olive-gray with small dark dots. venter smooth.5 foot. middorsal stripe beginning on the snout and continuing to occiput. a wide slate chevron mark above the shoulders dividing posteriorly into two dark stripes enclosing a light central area on the lumbar region and widening and becoming irregular on the sacral region. hand. upward and around arm insertion to mark a pale semicircle around the axilla. This difference may result from age and manner of preservation. when hind legs are bent at right angles to body.5 mm. ending above shoulder. Its back is smooth to pustular. mm. with a few very weak granules on the sacrum. head length.5 and body. rounded outer one. especially concentrated below a light diagonal line entering the groin but not extending forward beyond the level of the adpressed elbow. venter immaculate pale cream-buff. soles of feet and palms of hands olive-gray. in postaxillary region a little narrower than greatest When hind leg is adpressed. femur. no skinfold across the chest. heel reaches tip of snout. a heavy tarsal ridge. side of head olivegray. heels barely touch. not far from the type — 337-262—70 6 . as do the differences in size of finger and toe disks. wide glandular ridge from posterior corner of eye above tympanum. may represent the dark middorsal head stripe of the former. 7. a distinct skinfold on heel and knee. width of head. a weak oval inner metatarsal tubercle and a more distinct.. a slight ventral disk. except in a few points. and the disks slate-gray. with a slate-black stripe from nostril to eye through loreal region. Two examples from Pueblo Nuevo near Nilo. no exlaid along the sides. with some very weak tubercles on sacrum. 6 — Head 7. posterior femur with a pearl gray irregular stripe limited ing above by the ends of the crossbars and below by some scattered webs slate-gray spots. tibia. mm. This young specimen from the type locality agrees reasonably well with the type description. Remarks. ternal vocal sacs in this specimen.FROGS OF COLOMBIA —COCHRAN AND GOIN 69 penultimate phalanx of fourth. disk of fourth toe covering about one-third the tympanum and about the same size as that of third finger.. Body moderately slender. continuing behind eye on wide supratympanic ridge and curvColor in alcohol. an indistinct... 11 mm. Dimensions.5 mm. head width. Skin of upper parts resistant to abrasion. arms and legs very light drab-gray with three (on tibia) or four (on femur) wide slate crossbars. the between the toes white. blackish slate.. while the type has the back finely granular. — Back olive-gray with a wide.5 mm. mm.

205. 453. pi. pp. 19. locality.— Kellogg. p.— Taylor and Smith. 1930. 537. Rana palmipes Spix. 64 (type 13. 1952. 537. 1959.— Peters. Peru. disks above or below.— L. 93. 284. 202. p. MLS 309.— Smith and Taylor. 37. Rana palmipes Spix Plate 12a-c 1824. and very faint in the third. 473. 420. 165. 412.— Lutz and Kloss. p. 1931. p. p. 415.— Breder. 908. 1946. 91. 1888a. 1927. Ribeiro. the adpressed heel reaches between the eye and the nostril. 802. p. 23. 22. p. 1932.— Crawford. 622. p. 5. Lake of Junin.— Mirandapis. p.— Baumann. 92. 1961. p. p. 151876-7.— Cope. 1920.— Mello-Leitao. 521. or T-shaped. 1873c. 210 (type species. 1937. 1891a. fig. 1882a. 676. 113. p. In one example. (type locality. 1927. p. temporaria). 1958.— Crawford and Jones. Genus Rana Linnaeus 1758. p. p.— A. 1932. notched behind. — Pupil horizontal. USNM southwest of Fusagasuga. Rana juninensis Tschudi. 98. p. Tympanum distinct or hidden. pi. 1952. "Amazonenfluss"). agree well type.— Boulenger. 1873b. Stebbins and Hendrickson. 282. 1959. 1026. 1918. 176. .— Goin and Layne.— Schmidt. 505. 1923b. 1951. 1945. p.. fig. 14. 1963a. Pueblo Nuevo. p. p. The venter is immaculate with locality. 57. Subfamily Raninae Sternum bony. p. 1952. 1948. 56. p. Miiller. 449.). Lutz. 1939. p. p. 1935. Two km. p.— Johnson. 1964b.— Taylor. 1845. figs. 1887b. 622. Rana Linnaeus.S. 91.— Aleman. 5. 161 et seq— Noble.— Roux. Tongue free and deeply Generic diagnosis. The dark bars across the femur are sharp in one example. 9. 605. Vomerine teeth present.— Schubart. p. 1944c. p. 292. Gines. 1937a. 1919. p. 1926. 1939. figs. p. 1901. p. I Peters.— Gorham. 1920. Outer metatarsals separated by a web. 1 map. 48.— Giinther. 1912. moderate in another. Omosternum and sternum with a strong bony style. NATIONAL MUSEUM BULLETIN 2 88 MLS 309 and with the description of the more than to a little less than one-half webbed. R.— Hellmich. otherwise. 4. Specimens Examined COLOMBIA Ctjndinamarca: near Nilo. 7-9. p. p. 161. p. 1913. p. p. p. p.— NicSforo Maria. Terminal phalanges acute. p. 28. 278. p. 1952. p. 2 1926. pi. 316.— 70 U. 210. p. p. 51. transversely dilated. 473. p. p. the pattern is similar in all three. 200. 1933. p. p. 1873a. Terminal without digits pointed or slightly dilated. p. — — 1963.000 ft. p. p. 26.— Schmidt and Inger. 1919. 40. 40. 140. 311. figs.— Dunn.— Parker.— Beebe.— Rivero.— Boulenger. The toes in these three specimens are a little except for some dark suffusions on the chin and throat.

p. 400 m. 1920. 1945. 1-lc— Boulenger. p. p. equal to interval between nostrils. 1882. Ecuador). p. p. 1871). Vomerine teeth in two small. Jericho (3. pi. 166. 201. 473. p.— Cope. 403. Rana melanosoma Giinther. 1871. Rana clamata guianensis British affinis Peters. fig. p. 161. 1920. 1920. (type locality. 205. Venezuela) p. — Baumann. —Boulenger. p. p. slanting. 1920. 1871b. p. preoccupied by Peters. b (type localities. 473. 1877. — Boulenger. p. Peru). sloping sharply to upper lip. 9. 373. 1920. p. p. Brocchi. 1882a. Ranula affinis. Colombia. 117. p. ovate. p. pi.— Boulenger. 1920. 131 (type locality. 1871. FROGS OF COLOMBIA —COCHRAN AND GOIN 71 1859. Boulenger. —Steindachner. Boulenger. 402 (type locality. its diameter equal to its distance from tip of snout. 1874. Ranula brevipalmala Cope. Nauta. 1929. rounded at the tip when viewed from above. 175. p. — Cope. Rana vaillanti Brocchi. Bucay. p. 473. — — USNM Description 142198. Rio Napo. the upper jaw extending somewhat beyond the lower. 49. p. 1866b. 1867. 1867. Hylarana brevipalmala. b (type locality. 473. Rana nigrilatus. widely separated series between the choanae. Nicaragua). Guiana). p. 412 Peters. Tympanum distinct. Ranula palmipes. 473. 1872. 1920. interorbital diameter slightly less than that of upper eyelid. pi. 1920. pi. p. British Honduras). p.Barbour and Loveridge. Island. 1877. 130. 131 (type locality. Rana bonaccana Giinther. 15.— Boul- p.— Boulenger.— Boulenger. p. 49 (substitute for Ranula [ Rana] brevipalmata Cope. an adult from Leticia. 1882. their distance from end of snout a trifle less than their distance from eye. enger. 60. p. 473. 161.— . 1882. 1920. 1870. loreal region concave. — Boulenger. 473. 1874b.). p. p. p. p. Boulenger. brevipalmala rhoadsi Fowler. 1901. 1912. p. 166. Ranula gollmcri Peters. p. prominent. 205. 326. 1920. 222. — — 1920. 203.— Boulenger. 473. 1913. p. Rana palmipes rionapensis Andersson. p. p. Province of Guayas. Duenas. 1920. and Hacienda Rosa de 1913. Eye large.— Boulenger. off the coast of British Honduras).) — — — 1920. 1874. Canthus rostralis straight. Rana 1859. fig. 1882. 1877. 1. p. 473. rounded in profile. p. 473. 403 (type locality. Rana Rana 1913. p. figs. 1901. Bonacca — Boulenger. 473. 1863. 1912. 11. brevipalmala. 63. p. Caracas. . 1866. Rana copii Boulenger. Nostrils dorsolateral. Caracas. Venezuela). p. 1868. separated from eye by an interval equal to one-fourth . p. 1863a. pi. 62. p. (type locality. 1920. 1920. Guatemala. Peru). projecting. Amazonas. 473. upper fig. Mullins River near Belize. Boulenger. 3 (type locality. 1920.250 ft. its greatest width twothird that of eye. 473. Ranula nigritatus. 1872. 473. p. = Baumann. 167. 175 (type locality. snout elongate. 114. 1874. p. Pohlia palmipes. figs. 9. prominent. 1882a. Nauta. 5-8. 2.. 1920. Boulenger. p. Fowler. Ranula nigrilatus Cope. pi. tongue two-third as wide as mouth opening. 473. 1876. Ecuador. its posterior border free and very deeply notched.

metacarpal tubercles well developed. a small elongate inner metatarsal tubercle no outer tubercle a heavy skinfold on heel and knee. a pronounced ventral disk. Colombian specimens at hand are from Aruazonas. their tips reaching almost to base of antepenultimate phalanx of fourth toe. when limbs are laid along the sides. external vocal sacs. with than second and equal to both second and fourth reaching midway on penultimate phalanx of third.5 mm. When hind leg is adpressed. a wide brown stripe slightly broken by buff spots along forearm from base of thumb to elbow.72 its U..5 mm. a heavy glandular ridge from posterior corner of eye above tympanum. head length. webs of toes mottled with brown and buff. tympanum nearly encircled by a brown ring. ending behind and below corner of mouth a distinct skinfold across the chest. Meta. own NATIONAL MUSEUM BULLETIN 288 diameter. in postaxillary region less than greatest width of head. with a few faint brown spots on chin and chest. Santander. a small rounded palmar callus. 35 mm. 28. free. hand. Antioquia.. tibia. paler below and heavily spotted with dark brown. venter pale buff. Color in alcohol —Dorsum smoke gray with irregular drab motthngs. heel reaches to anterior corner of eye. when hind legs are bent at right angles to body. 21. Toes entirely webbed.. a pair of glandular dorsolateral ridges from posterior corner of eye almost to groin. 30.5 mm. Body moderately slender. the tips of . upper parts of legs alternating with — . and possibly demand the description of still other forms. compiled mostly from Boulenger's and Taylor's writings. and feet -with three wide transverse brown bars two or three paler ones. first finger longer fourth. an elongate thumb pad present. Skin of upper parts smooth. heels overlap slightly. the above synonymy. Cauca. Fingers moderately long. posterior femur coarsely marbled with clove brown and pale buff. foot 37. Choc6. is more or less accepted at the present time. side of head drab gray. except for numerous small transverse wrinkles from level of axilla to groin. the lower sides of toes and tarsus quite dark. with a narrow darker gray line indistinctly paralleling the edges of both upper and lower jaws. Cundinamarca. . head width. Caqueta.S. lower surfaces of legs buff with many small dark round spots. apparently no distinct lateral ridges.. 75 mm. the tips slender.5 mm. knees and elbows considerably overlap. third and fifth subequal. femur. anterior femur with a few brown spots on the pale buff ground color. Tolima. a few small pustules on sacral region and above vent. sides smoke gray to drab above. 38 mm... venter smooth. — Dimensions Head and body. . Remarks Although a thorough comparison of all available material throughout the vast range of this frog may validate some names now considered as synonyms.

on Rio Ariari.. a careful study of all available Rana palmipes from throughout its range is needed to the amount of geographical variation. Sierra de La Macarena. the tadpole is still in metamorphosis. 6384S.. 10-15 km. 22599. 144538-9. near El Refugio. . 185 m. USNM 144537.). 146253-4. ear. The measured (MLS from Villa vicencio. Andagoya. 39279. Medellin Valley. USNM USNM of Bucaramanga. — COCHRAN AND GOIN 73 species probably occurs also in other parts of the country. USNM 142198-205. 150522-4. Antioquia). 39275-6. USNM USNM Santandek: 4. and a scattering of large dark spots on the sacral region and center of back can be seen. 144543. Cundinamarca: Fusagasugd. USMN USNM 144544-53. Lagos de Cacique. MLS 266. Puerto Narifio. Tolima: USNM AMNH Mariquita. 147271. only Colombian examples are listed here. measuring 40 millimeters. upper Rio Giiejar and El Mico. 147123-4. Granada. and mouth were completely developed. so that the dark leg bands stand out more clearly.140 m. Meta: Angostura. As such a study cannot material of discover be attempted in this paper. northeast of Bellavista. The large old specimen from Villavicencio (MLS 266) has the toe webs nearly uniform brown in color. In 39275 (42 millimeters. 39463-4. USNM 144534-6. legs. above Rio Porce. El Centro. Cafio del Morrocoy. 151875. tributary of Atrato River. USNM 144540. Colombian Specimens Examined Amazonas: USNM Leticia. largest specimen Meta) has a 266. Rio San Juan. USNM 146418-20. tributary of Rio Guayabero. 144554-5. Villavicencio. south of Medellin. 151473-5. 39258-64. Atrato River. USNM 151902. Near Moscopan. USNM USNM USNM 150525. the arms and legs had appeared and the tail was reduced to 15 millimeters in length. 1.200 m.. Some of the frogs from Leticia have a much paler ground color than the described specimen. 1. while the dark spots are concentrated along the sides and in the groin. AMNH AMNH As for other wide-ranging species. 144531. southeast 146249. Sonson. AMNH 13721. four km.FROGS OF COLOMBIA and The Valle. and also from Envigado). Chigorod6. Antioquia). Cauca: Choc6: USNM 147217-20 (juv. Mouth Caqueta: AMNH CNHM Rio Mecaya. of USNM 144532-3. In 39276 (from Envigado. Rio Mecaya. In a tadpole measuring 28 millimeters in head and body length (AMNH 22600 from Sonsdn. AMNH 13602. Cafio Guapaya. total length of 110 millimeters. south of Medellin. the arms. AMNH AMNH Antioquia: Envigado.7 km. west of Playa de Oro. 22600-3. Quesado River. CNHM 69750-1. Lebrija. with neither the adult tympanum nor mouth formed. Quebrada Honda. 39256. Boca de la Raspadura. 39266. 39273. 144541-2. while a very slight tail vestige remained.

Relicti vomer Posterior portion of prevomer present Posterior portion of prevomer absent Elachistocleis Clavicles absent. lower Rfo Calima. Glossostoma Ctenophryne Otophryne Palatines present Palatines absent Tympanum distinct Genus Relictivomer Carvalho 1954. 121.S. . — Parker. Hypopachus pearsei Ruthven). 1954.— Barbour and Loveridge. They have no ribs or intercalary cartilages. c2. p. Only one species is known from Colombia. Tympanum concealed. — Dunn. p. 1934a. c b2 to concealed. 149738-40. 151464-82. near C6rdoba. Elachistocleis pearsei. The larvae lack horny mandibles and labial teeth. 1954. and procoracoid present but short and curved. a tongue. 26. 1944. Fundaci6n. NATIONAL MUSEUM BULLETIN USNM USNM USNM Valle: Rio Calima. from Glossostoma aterrimum by the absence of the palatine bone. p. 145746-55. reduced to a small osseous plate which lies in the palatal region. — Gorham. Relictivomer pearsei (Ruthven) Figure 1914. 1922. 1946. 13. . Key a1 . 18. 77 (type locality. — A microhylid with the tympanum concealed. the posterior portion of the prevomer present. p. 1952. 50.— Ruthven. Palatine bones absent. Relictivomer Carvalho. p. 13 (type species. Colombia). p. p. Family 2 88 Camp Carton de Colom- USNM 151476. 1934. El Tigre. in which the pale of the belly is continuous with the light color in the From Otophryne robusta tympanum concealed. Clavicle present but short and curved. from axillary region so that there it is no discrete axillary pale spot. and from Elachistocleis ovalis by the presence of the posterior portion of the prevomer and by the lighter colored belly.74 U. may be distinguished by having the Ctenophryne geayi by the presence of the clavicle and procoracoid. bia. clavicle Diagnosis. 1944c. 1 Hypopachus pearsei Ruthven. Virology Microhylidae Frogs with firrnisternal girdles. and a very pointed snout. Rfo Raposo. are paired. — Prevomer divided with the posterior portion Generic diagnosis. 1963. p. snout short and very pointed. c2 . Elachistocleis ovalis (not of Schneider). and sacral diapophyses more or less expanded. and Eustachian tubes. if present. 132 — Peters. p. c a2 Colombian Genera of Microhylidae Clavicles present. Relictivomer pearsei. palatine bones absent. . 19. 524: 1949. p. 1 . Procoracoid present. Tympanum b 1 1 . Field Station. . — Carvalho.

paratype. .— FROGS OF COLOMBIA — COCHRAN AND GOIN 75 — USNM 51215 (paratype). separated from each other by an interval but slightly less than their distance from eye. Description. pointed when viewed from above. Eye small. Magmicrohylid without vomerine teeth. the upper jaw extending considerably beyond lower. inconspicuous. from Fundaci6n. their distance from end of snout about one-half that from eye. Snout short. pointed in profile. an elongate oval in form. not at all projecting. Figure 1. A Relictivomer pearsei. nostrils more lateral than superior. palpebral membrane not reticulate. Can thus rostralis very slightly defined. its posterior one-half free and unnotched. interorbital distance about three times the width of upper eyelid. loreal region rounded. choanae rather small and placed well forward in the roof of the mouth. tongue half as wide as mouth opening. Colombia. USNM 51215 (X 2). dalena. its diameter slightly less than its distance from nostrils.

12. and the groin and anterior and posterior surfaces of the thighs distinctly marbled in yellow and brown.S. — — Undersurfaces of throat. Body short. fourth about equal to second. reaching to base of penultimate phalanx of third. are rather dark with a single narrow. with a few brownish flecks under throat and on lower surfaces of thighs.7 mm. hand In this individual the belly. Our largest specimen has a head-body when sufficient material length of 41. Remarks. femur. in postaxillary region at least twice the greatest width of head. 7. 2 88 which is about two-thirds the distance between nostrils. and lower surface of thigh smooth. Fingers unwebbed. 12. no vocal sac. no dermal appendage on heel.8 mm.. when hind legs are bent at right angles to body. no projecting rudiment of a pollex. no ulnar ridge. species tend to ation in the Maracaibo Basin. heels overlap appreciably. Head and body.76 NATIONAL MUSEUM BULLETIN U. 33. 1954) saw . third toe two phalanges longer than fifth. head width. Color in alcohol. is nearly marbled as uniformly pale yellow. knee and elbow fail to meet. no tarsal ridge. Except for the specimen discussed above. Top of head and dorsum uniform warm brown.4 mm. Tympanum concealed. show some degree it is of incipient speci- not too surprising to find the single specimen of pearsei we have from there different from other individuals from west of the Cordillera Occidental. when hind leg is adpressed. Skin of throat. head length.. hand. chest. 8. 8. adult female. Variation. It may well be that becomes available the population in the Maracaibo Basin will prove to be deserving of a name. belly. tibia. there seems to be no significant variation in the specimens at hand. belly..6 mm. The posterior surfaces of the thighs. but whether this is due to preservation or individual variation we do not know.5 millimeters.8 mm. no apparent traces of a skinfold across chest. A narrow fold of skin crosses back of head and passes downward on each side behind eye and angle of jaw. — Since the recent monographer of this group in South fit to place this species in the genus America (Carvalho. on the other hand. Some are darker than the others. instead of being conspicuously and brightly it is in other specimens of Relictivomer pearsei.. Dimensions. chest. heel-to-toe. and limbs. is —The most notable exemplified by the specimen variation in the specimens at (MLS 133) from Astillero. Skin of upper parts smooth. Structurally this specimen does not seem to be different from other examples of pearsei in Since so many any way. yellow stripe running from behind the knee to the vent on each of them.3 mm. when limbs are laid along the side. 20 mm. a distinct but small oval inner but no apparent outer metatarsal tubercle... Toes without webs. heel reaches axilla.

FROGS OF COLOMBIA

— COCHRAN

AND GOIN

77

and since we have had very little first-hand experience
with the other genera in South America, we follow him in the generic
assignment. We must say, however, that, to us, ovalis and pearsei seem
to be rather closely related, and in fact we must agree with Dunn
(1949, p. 13) "I would take them for vicarious races save for the
statement of Ruthven that the two occur together at Fundaci6n."

Relictivomer

:

Specimens Examined

COLOMBIA

AMNH 13539.
CNHM

Antioquia: Medellin,
Choc6: Sautata, Rio Atrato,

7490S.

Magdalena: Fundaci6n, USNM 51215
Norte de Santander: Astillero, MLS

(paratype).
133.

Santander: El Centro, 112 m., AMNH 71044.
Tolima: Mariquita, MLS 134, CNHM 81817.
No Precise Locality: USNM 14718.
PANAMA: Chiriqui Province: 0.7 mi. north and
69876-7.

Panama

Province:

3 mi. south of Bejuco,

0.2 mi. west of David,

Nueva Gorgona,

mi. west of

AMNH

69880-3;

Elachistocleis Parker

Elachistocleis Parker, 1927a, p. 4 (type species,

Generic diagnosis.

AMNH

AMNH 69878-9.

Genus
1927.

1

— Posterior portion

of

Rana

ovalis Schneider).

prevomer absent. Palatine

bones absent. Clavicle present but short and curved. Procoracoid
present. Tympanum concealed, snout short and very pointed.
Only one species is known from Colombia.
Elachistocleis ovalis (Schneider)

Figure 2
1799.

Rana

ovalis

Schneider, p. 131 (type locality not given).

— Shaw,

1802,

p. 111.

1802.
1802.

1820.
1826.

Bufo surinamensis Daudin, p. 91 (type locality, Suriname) 1803, p. 184.
Bufo ovalis.— Daudin, 1802, p. 92; 1803, p. 187.
Rana bufonia Merrem, p. 177 (substitute name for Bufo surinamensis
;

Daudin).
Engystoma ovale. Fitzinger, 1826, p. 65. Dumeril and Bibron, 1841, p.
741.— Gunther, 1858, p. 51.— Steindachner, 1864, p. 285.— Hensel,

140.— Boulenger, 1882a, p. 163; 1886a, p. 439; 1894, p. 347;
1898b, p. 131.— Cope, 1885b, p. 185; 1887, p. 53.— Boettger, 1885, p.
240; 1892, p. 22.— Peracca, 1895, p. 23; 1904, p. 11; 1914, p. 104.— Berg,
1896, p. 159.— Werner, 1899, p. 475.— Budgett, 1899, p. 305.— Lidth,
1867, p.

1904, p.

1924, p.
1830.

93— Mehely,

1904, p. 211.— Baumann, 1912, p. 129.— Marelli,
585.— Miranda-Ribeiro, 1926, p. 184.— Lutz, 1927, pi. 8, fig. 2;

1928, p. 92.
unicolor

Microps

Wagler,

p.

200

(substitute

name

Schneider).
1910.

Gastrophryne

1920.

Engystoma

ovalis.

— Stejneger, 1910, 166.
— Miranda-Ribeiro, 1920a,

ovale ovale.

p.

p. 282.

for

Rana

ovalis

.

78

U.S.

NATIONAL MUSEUM BULLETIN

2

88



1926.

Engystoma ovale concolor. Miranda-Ribeiro, 1920a, p. 284 (type locality,
Ypiranga) 1926, p. 184.
Engystoma ovale lineala. Miranda-Ribeiro, 1920a, p. 2S4 (type locality,
R. da Serra, Sao Paulo).
Engystoma ovale caesarii. Miranda-Ribeiro, 1920a, p. 284 (type localities,
Piquete, Os Perus, Alto da Serra [Sao Paulo], Cubatao [Santos], and
Ypiranga)
Engystoma caesarii mottae Miranda-Ribeiro (nom. nud.), 1920a, p. 284
(type locality, Sao Paulo), 1926, p. 194.
Gastrophryne ovale,
Ruthven, 1922, p. 51.
Engystoma ovalis. Nieden, 1926, p. 66.

1927.

Elachistocleis ovale ovale.

1929.

286.— Dunn, 1931, p. 416.
Elachistocleis ovalis.
Mertens, 1930, p. 163.— Crawford, 1931, p. 38.
Parker, 1934, p. 121.— Myers, 1942, p. 155.— Dunn, 1944, p. 524; 1949,
p. 13.— Carvalho, 1954, p. 15.— Rivero, 1961, p. 177; 1964a, p. 304.—
Gorham, 1963, p. 26.

1920.

;

1920.
1920.

1920.
1922.

1930.


— Parker,

1927a, p.

4.

Elachistocleis ovale.— Mertens, 1929, p.

Diagnosis.

—A microhylid with the tympanum concealed,

clavicles

and procoracoids present but short and curved, posterior portion of
the prevomer absent, palatine bones absent, and snout very pointed.
The presence of clavicles and procoracoids separates Elachistocleis
ovalis from species in the genera Ctenophryne and Glossostoma, while
the concealed tympanum and the fact that the clavicles and procoracoids are short and strongly curved separate it from species of
Otophryne. From Relictivomer pearsei it may be distinguished by having
the belly dark with isolated light spots on the anterior face of the

Figure

2.

Elachistocleis ovalis,

FM

266 (same

size).

;

FROGS OF COLOMBIA

— COCHRAN

AND GOIN

79

it joins the body and usually present in the axillary
and by having the posterior portion of the pre vomer absent.
FM 266, from Villa vicencio, Meta, Colombia. A
Description.
microhylid frog without vomerine teeth; choanae moderate in size,
ovate, placed well forward in the roof of the mouth. Tongue half as
wide as mouth opening, an elongate oval in outline, its posterior half
free and unnotched. Snout short, pointed when viewed from above,
pointed in profile, the upper jaw extending considerably beyond lower;

thigh where

region,

more

than superior, not at all projecting, their distance
about one-half that from eye, separated from each
other by an interval equal to about two-thuds their distance from eye.
Canthus rostralis but very slightly defined; loreal region distinctly
rounded. Eye small, inconspicuous, its diameter less than its distance
from nostril; palpebral membrane not reticulate; interorbital distance
about three times the width of upper eyelid, which is even less than
distance between nostrils. Tympanum concealed. Fingers unwebbed,
fourth finger about equal to second, reaching to base of penultimate
phalanx of third; no projecting rudiment of a pollex; no ulnar ridge.
Toes without webs, fourth toe nearly two phalages longer than third
a distinct oval inner but no apparent outer metatarsal tubercle; no
tarsal ridge; no dermal appendage on heel. Body short, in postaxillary region at least twice the greatest width of head; when hind
leg is adpressed, heel reaches axilla, when limbs are laid along the
side, knee and elbow fail to meet; when hind legs are bent at right
angles to body, heels touch snugly. Skin of upper parts smooth. A
narrow but distinct fold of skin crosses the back of the head and turns
down on each side to pass behing the eye and the angle of the jaw.
Skin oi throat, chest, belly, and lower surface of thigh uniformly
smooth; adult female, no vocal sac.
Dimensions. Head and body, 43.5 mm.; head length, 10.1 mm.;
head width, 11.1 mm.; femur, 15.8 mm.; tibia, 15.9 mm.; heel-to-toe,
18.4 mm.; hand 10.8 nun.
Color in alcohol.- -Top of head and dorsum dark grayish brown.
Underside of throat, chest, belly, and limbs faintly flecked yellowish
gray. Rather distinct yellowish subaxillary and groin spots and a
narrow yellowish line on the posterior surfaces of the thighs.
Variation.
The specimens at hand, all from the northern part of
the range of the species, seem to be rather uniform in both pattern
nostrils

from end

lateral

of snout

and

structure.

The

rather distinctive, isolated yellow spot in the

and the one on the anterior face of the thigh where it meets the
groin apparently are quite good key characters, for the inguinal spot
on the thigh is present in every individual and the axillary spot
appears in all except one specimen from Villa vicencio. These same
yellow areas are present in 11. pearsei, but in that species such areas
axilla


80

NATIONAL MUSEUM BULLETIN 288

U.S.

do not form distinctive isolated yellow spots, being continuous with
the yellow of the belly. The largest specimen of Elachistocleis oualis
at hand has a head-body length of 39.3 millimeters.
Specimens Examined

COLOMBIA
BoyacX: Corozal, near Rio Casanare, USNM 152032-3.
C6rdoba: Junguilla, Cano Betancf, USNM 152158.
Meta: Cano Guapaya, Serrania de La Macarena, CJG 2340;

CNHM 81816, USNM
USNM 151967-76.

152204-8,

MLS

132, 132a;

Villavicencio,

Granada, on Rio

Ariari,

USNM 152316.
USNM 128847-9.

Vichada: Amenavn6, Rio Guaviare,

VENEZUELA:

Palenque, Guarico,

Genus Glossostoma Giinther
1900.

Glossostoma Giinther, p. 210 (type species, Glossostoma aterrimum Giinther) #

Generic diagnosis.
present.

—Clavicles and procoracoids absent, but palatines

Tympanum

Only one

species

is

concealed. Snout not particularly pointed.

known from Colombia.

Glossostoma aterrimum Giinther

Figure

3

Giinther, p. 210 (type locality, Costa Rica).
Boulenger, 1913, p. 1025.— Nieden, 1926, p. 68.— Carvalho, 1954, p.

1901.

Glossostoma aterrimum

1927.

Gastrophryne aterrimum. Parker, 1927a, p. 5.
Microhyla aterrimum. Parker, 1934a, p. 143.
1949, p. 16.— Taylor, 1952, p. 914.

11.— Gorham,
1934.

Diagnosis.

and

1963, p. 26.

—A microhylid with

the

— Dunn,

tympanum

1944c, p. 524.

concealed, clavicles

procoracoids absent, but palatines present.

Glossostoma aterrimum may be distinguished from species of the
genera Relictivomer, Elachistocleis, and Otophryne by the absence of
clavicles and procoracoids and from those of Ctenophryne by the
presence of palatine bones.
Description.
63849, four kilometers northeast of Bellavista, above Rio Porce, Antioquia, Colombia. A microhylid without
vomerine teeth; choanae rather large and broadly rounded; tongue
two-thirds as wide as mouth opening, broadly ovate, its posterior
border free and unnotched. Snout large for a microhylid, rather rounded
when viewed from above, nearly truncate in profile, the upper jaw extending slightly beyond lower; nostrils more lateral than superior,
very slightly projecting, their distance from end of snout about
two-thirds that from eye, separated from each other by an interval
slightly greater than distance from eye. Canthus rostralis poorly

— CNHM


FROGS OF COLOMBIA

—COCHRAN

defined; loreal region very slightly concave

AND GOIN

and nearly

81
vertical, the

upper lip flaring out but little below it. Eye small, not prominent, its
diameter equal to its distance from nostril; palpebral membrane not
reticulate; interorbital distance about one-third the width of upper
eyelid, which is relatively narrow and distinctly less than distance
between nostrils. Tympanum practically concealed, just barely discernible at the surface on the right side, concealed on the left. Fingers
without a trace of web at base, fourth finger a phalanx longer than
second, just reaching to base of terminal phalanx of third, tips without
digital expansions; no projecting rudiment of a pollex; no ulnar ridge.
Toes slightly more than one-half webbed, the web on fourth toe reaching the base of the antepenultimate phalanx, third toe much longer
than fifth; no metatarsal tubercle apparent; no tarsal ridge; no dermal
appendage on heel. Body rather elongate, in postaxillary region a
little wider than greatest width of head; when hind leg is adpressed,
heel reaches almost to angle of jaw; when limbs are laid along the side,
knee and elbow meet; when hind legs are bent at right angles to body,

Figure

3.

Glossosloma aterrimum,

CNHM 63849

(same

size).

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U.S.

2

88

heels overlap slightly. Skin of upper parts smooth; a narrow, indistinct
skinfold across back of head which, behind the eyes, becomes more

and passes downward and backward to terminate behind the
body smooth except under the throat
where it bears distinct, rounded, pimple-like granules and the lower
surface of the thigh which is uniformly and very finely granular;
no traces of a skinfold across chest; adult male; no vocal sac apparent.
Dimensions. Head and body, 62.2 mm.; head length, 15.2 mm.;
head width, 20.5 mm. femur, 27 mm. tibia, 29 mm. heel-to-toe, 41.7
distinct

angles of the jaw. Skin of

;

;

;

mm.

mm.; hand, 16.8

Color in alcohol.

—In preservative the specimen

is

rather uniformly

medium dark brown above, pale brown below. No evidence
distinctive pattern

Remarks.

of

a

discernible.

is

—We have had the opportunity

of

examining the specimen

(1949, p. 16) reported from Muzo and of comparing it with the one described above, and for the present, we cannot

(MLS 247)

that

Dunn

agree with Dr.

Dunn

that

it

is

surely Glossostoma aterrimum.

The

webbing on the feet is only about half as extensive as on the specimen
described, and we cannot even palpate an unquestionable palatine.
This latter, however, may be due to the condition of the specimen,
for it is very soft; in fact, no hardened bones are palpable in any part
of the suspensorium. In color and general build the two specimens
are quite similar.
G. aterrimum is a distinctly web-footed microhylid from west of the
Cordillera Oriental of Colombia, an analogue of Ctenophryne geayi
east of the Cordillera. While both species have webbed feet, we suspect

evidence of convergence rather than close kinship, for the
different in general appearance, and the fact that palatines are present in Glossostoma and absent in Ctenophryne does not

that this

is

two are quite

seem indicative

closely related

of

species.

Specimens Examined

COLOMBIA:
Antioquia: 4 km. northeast of Bellavista, above Rio Porce,
Botaca: Muzo, MLS 247.
Caldas: Palestina, USNM 152145.

CNHM 63849.

Genus Ctenophryne Mocquard
1904.

Ctenophryne Mocquard,

Generic diagnosis.

Tympanum
Only one

p.

308 (type species, Ctenophryne geayi Mocquard)

—Clavicles,

procoracoids, and palatines absent.

concealed. Snout pointed.
species

is

known from Colombia.


FROGS OF COLOMBIA

— COCHRAN

AND COIN

83

Ctenophryne geayi Mocquard
Figure 4
1904.

Ctenophryne geayi Mocquard, p. 308 (type locality, Sarare River, Colombia).— Nieden, 1926, p. 69.— Parker, 1934a, p. 123.— Dunn, 1944c,
p. 524; 1949, p. 18.— Guib6, 1949, p. 59.— Carvalho, 1954, p. 7

Gorham,
Diagnosis.

1963, p. 26.

—A

microhylid

having

a

concealed

tympanum and

lacking clavicles, procoracoids, and palatines.

The absence
while the

and procoracoids separates this species
and Elachistocleis ovalis,
the palatines separates it from Glossostoma

of the clavicles

from Relictivomer

pearsei, Otophryne robusta,

absence of

aterrimum.

—MHNP

03-84 (holotype), from Rio Sarare, Norte de
Description.
Santander, Colombia. A microhylid without vomerine teeth; small,
rounded choanae; tongue three-fourths as wide as mouth opening,
pointed anteriorly, with well-developed cornua on each angle posteriorly, the cornua being free, a transverse row of glandular structures
across roof of mouth.

Snout small, triangular when viewed from

above, broadly pointed in profile, the upper jaw extending beyond

more lateral than superior, not projecting, their distance from end of snout about two-thirds that from eye, separated
from each other by an interval equal to their distance from eye.
Can thus rostralis hardly defined; loreal region concave and more
lower; nostrils

flattened than oblique, the upper lip flaring out, then rolling in below

Figure

4.

Ctenophryne geayi,

FM

357 (same

size).

84
it.

U.S.

Eye very

from

NATIONAL MUSEUM BULLETIN

2

88

small, not prominent, its diameter equal to its distance

nostril; interorbital distance nearly twice internarial distance.

No

exposed tympanum; fingers without a trace of web, fourth finger
than second; no projecting rudiment of a poll ex. Toes
about one-half webbed, the web on fourth toe reaching the base of
the antepenultimate phalanx, third toe much longer than fifth; an
oval inner but no apparent outer metatarsal tubercle; no tarsal ridge;
no dermal appendage on heel. Body stubby, in postaxillary region
probably twice width of head when hind leg is adpressed, heel reaches
to anterior part of shoulder region; when limbs are laid along the
side, knee and elbow fail to meet; when hind legs are bent at right
angles to body, heels fail to meet. Skin of upper parts quite smooth,
a transverse skinfold across back of head which continues downward
over shoulder region to inner side of elbow; skin of throat and chest
smooth, that of belly and lower femur also smooth; no skinfold
across the chest; no inguinal gland; no vocal sac apparent.
Dimensions. Head and body, 43.3 mm.; head length, 12 mm.;
interorbital width, 6.6 mm.; femur, 17.6 mm.; tibia, 16.7 mm.; foot,
24.7 mm.; hand, 11 (±) mm.
Color in alcohol.
Pale brown above, a much darker brown below,
with a sharp line of demarcation between the two. The demarcation
starts at the nostril, passes through the eye and just behind the fold
to the inner side of the elbow, thence out along the dorsal surface
of the forearm to the inner side of the third finger; the tops of third
slightly longer

;

and fourth

have the dorsal ground color. The line of demarcafrom the outer side of the little finger around the

fingers

tion then runs

elbow to the axilla, along the side to the groin, along the face of the
thigh and shank, thence along the foot to the base of the fifth toe;
it runs through the vent, along the posterior surface of
the thigh to the inner side of the knee, thence to the ankle and along
the margin of the fifth toe to its tip. Thus the dorsal surface of the

posteriorly

the only one with dorsal ground color. Above, the only
a very thin pale hairline running from tip of snout to vent.
Below, the entire ventral surface is flecked with pale spots, these

fifth toe is

pattern

being a

is

little

larger

on the breast. The dark ground color on the
is heavily marbled with

inner face of shank and inner face of ankle
white.

Remarks.

—Specimen

FM

357 from Serrania de

confirms that Ctenopkryne geayi

is

in reality a

La Macarena

Colombian

species

and

adds additional credibility to the type locality as cited by Mocquard.
Dunn (1949, p. 18) has pointed out that the Rio Sarare actually
arises on the southeast slope of Paramo de Tama in Norte de Santander
and flows for about eight miles through Colombian territory before
it crosses into Venezuela. It is Dunn's supposition that the type


FROGS OF COLOMBIA

COCHRAN AND GOIN

85

was collected by a member (French?) of the boundary commission,
which certainly visited the spot where the Sarare crosses the border
and there established the altitude (640 m.).
The few specimens available do not throw much light on individual
variation, but they do indicate that the species is rather widespread.
The absence of the clavicles and procoracoids and the webbing on
the toes set this species off rather sharply from all Colombian microhylids except Glossostoma aterrimum. Thus in Colombia there is only
one web-footed microhylid east of the Cordillera Oriental of the
Andes (Ctenophryne geayi) and one west of it (Glossostoma aterrimum).
The presence of the palatine bones in Glossostoma and the quite
different habitus of the two genera make them readily separable.
The presence of webbed feet, a rather pale brown dorsum with a
middorsal pale hairline, and a venter of a brown darker than the
dorsum and separated from it by a rather sharp line of demarcation
seem to be rather uniform traits of the species.
Specimens Examined

COLOMBIA
Norte de Santander: Rio Sarare, MHNP 03-84
Meta: Serrania de La Macarena, FM 357.
BRAZIL: Amazonas, region of Rio Livranents,

(holotype).

AMNH 44787; Manjuru
AMNH (unnumbered specimen).
AMNH 23119, 71399, USNM 129533.
AMNH 42887; Peru-Brazil frontier (UtaquiniaPampa Hermosa, Rio Cushabatay, 500
AMNH

Mau6s, middle Amazon),
GUIANA: Kartabo,

(region of

BRITISH
PERU: Inguitos,
Tapiche),
42663.

AMNH

350 ft.,
42685;

ft.,

Genus Otophryne Boulenger
1900.

Otophryne Boulenger,

Generic diagnosis.

p.

—Clavicle

straight. Palatines absent.

Only one

species

55 (type species, Otophryne robusta Boulenger).

is

and procoracoid present and nearly

Tympanum

distinct.

known from Colombia.
Otophryne robusta Boulenger
Figure

1900.

5

Otophryne robusta Boulenger, p. 55 (type locality, Mt. Roraima, British
Guiana).— Nieden, 1926, p. 73.— Noble, 1931, p. 536.— Parker, 1934a,
p. 108.— Dunn, 1949, p. 2.— Carvalho, 1954, p. 3.— Rivero, 1961, p.

179.— Gorham,
Diagnosis.

1963, p. 26.

— Tympanum distinct; clavicle and procoracoid

and perpendicular to midline

Otophryne robusta can be easily distinguished from

now known from Colombia by
clavicles

straight

of pectoral girdle; palatine absent.

its distinct

all

Microhylid ae

tympanum and

the straight

and procoracoids which are perpendicular to the midline of

the pectoral girdle. All other Colombian species have the
337-262—70

7

tympanum


86

U.S.

NATIONAL MUSEUM BULLETIN 288

concealed; and the clavicles and procoracoids,

curved.

if

present, are strongly

—FM

89, from lower Rio Apaporis, Amazonas, Colomvomerine teeth; choanae moderate in size,
microhylid
without
bia. A
forward in the mouth; tongue not more
and
lying
well
ovate
laterally
as
mouth
opening, elongate, its posterior border
wide
than half as

Description.

and unnotched. Snout rather large for a microhylid, somewhat
when viewed from above, quite pointed in profile, the upper
jaw extending very considerably beyond lower; nostrils lateral, not
visible from dorsal view, not projecting, their distance from end of
snout slightly more than one-half that from eye, separated from each
other by an interval equal to their distance from eye. Canthus rostralis
sharply denned; loreal region flattened and flaring inward toward
the upper lip. Eye rather large for a microhylid, moderately prominent, its diameter equal to its distance from nostril; palpebral memfree

truncate

brane not reticulate; interorbital distance about three times the
width of upper eyelid, which is even less than the distance between
nostrils. Tympanum very distinct, its diameter about equal to
greatest diameter of eye, separated from eye by a distance less than
one-fourth its own diameter. Fingers without a trace of web at base,
fourth finger but slightly longer than second, just reaching to base

Figure

5.

Otophryne robusta,

FM

89 (same

size).

FROGS OF COLOMBIA

—COCHRAN

AND GOIN

87

phalanx of third; digital tips not expanded; no prorudiment of a pollex; no ulnar ridge. Toes slightly webbed
at base; the web on fourth toe reaching the base of the antepenultimate
phalanx, third toe much longer than fifth; metatarsal tubercles not
present; no tarsal ridge; no dermal appendage on heel. Body rather
elongate, in postaxillary region slightly broader than greatest width
of penultimate

jecting

head; when hind leg is adpressed, heel reaches axillary region;
when limbs are laid along the side, knee and elbow fail to meet;
when hind legs are bent at right angles to body, heels fail to meet
by internarial distance. Skin of upper parts smooth. A narrow dorsoof

from above the tympanum to the region of the
A rather narrow glandular ridge encircles upper
part of tympanum. Skin of throat, chest, and belly smooth; no
trace of a skinfold across chest; adult female, no vocal sac. Abdomen
packed with large, unpigmented eggs.
Dimensions. Head and body 51.7 mm.; head length 17 mm.; head

lateral fold passes

groin on each side.

width, 18.3

mm.; femur,

mm.; hand,

11.5

19.1

mm.;

tibia,

11.8

mm.;

heel-to-toe, 25

mm.

Dorsum uniform dark brown medially. A pale,
Color in alcohol.
narrow, cream-colored stripe extends from eye to eye around the tip of
snout, passing just above the nostrils. This stripe widens appreciably
just behind each eye and passes posteriorly on each side above the
tympanum in the form of a broad, cream-colored dorsolateral stripe
all the way to the groin, where it again narrows and passes along the
anterior faces of the thigh and shank as a narrow cream-colored stripe.
Posterior side of thigh dark with light mottlings arranged

more

or less

form of a stripe. Lower surfaces somewhat paler than dorsum,
darkest under the chin and throat, becoming paler posteriorly. Indistinct but discrete, rounded, scattered, brown ocelli-like dots on the
chin, throat, and chest.
Remarks. This very distinctive genus is still, to the best of our
knowledge, known from only four specimens the type; a specimen in
the American Museum (AMNH 1325) which has no data other than
that it was obtained from the museum in Georgetown, British Guiana;
a specimen in the University of Michigan collection (MZUM 85137)
from Arabop6, Venezuela; and the specimen, described above, from
the lower Rio Apaporis in Colombia.
This is surely the most distinctive microhylid in South America.
The large, conspicuous tympanum and the very flat-topped head set it
off at a glance from all others.
The specimen at hand, a gravid female, has the abdomen packed
with large, unpigmented eggs, a condition usually associated with
in the

terrestrial

We

breeding habits.

have not examined the type of

this species,

but from available

88

U.S.

NATIONAL MUSEUM BULLETIN 288

hand differs in three
and dark on the
posterior face of the thighs, with light markings arranged in the form
of a narrow stripe. The dorsolateral stripes are quite distinct and the
way they pass through the groin and onto the anterior surfaces of the
thigh and shank is unique among the frogs known to us. The dorsolateral folds are narrow and sharp and, on gross examination, seem to

descriptions

we judge that our specimen

at

respects: dorsolateral stripes; dorsolateral folds;

lack the glandular structure such as one finds in Rana, for example,
but they are too uniform and symmetrical to permit us to believe that
they are merely artifacts of preservation. The type is described as
having the groin and hinder surfaces of the thighs orange with black
spots. In the specimen from Colombia, the groin is dark except for the
dorsolateral stripe that passes through it and the hind surface of the
thigh is likewise black except for the narrow stripe formed by confluent
light markings.

The specimen from Colombia has been rather grossly dissected and
we cannot now find any sign of an omosternum such as is illustrated
by Parker (1934a, fig. 46). The condition of the girdle, however, does
not permit us to state categorically that one was never present for a
small one such as Parker illustrates may well have been inadvertently
removed in the process of cleaning the girdle. Hence, like Dunn (1949)
and Carvalho (1954) we cannot confirm the presence of an omosternum
in this genus.

Specimens Examined

COLOMBIA
Amazonas: Lower Rfo Apaporis,

VENEZUELA:

Arabop6,

FM 89.

MZUM 85137

Suborder
Vertebral

column

(photograph).

PROCOELA

procoelous,

although

sometimes

free

inter-

vertebral disks are formed. Pectoral girdle usually arciferal, with
posteriorly projecting epicoracoid horns always present. Epicoracoids

sometimes more or less fused. Urostyle articulates with sacral vertebra
by a double condyle (sometimes urostyle and sacrum are fused).
Ribs never present.

Family Pseudidae

Each

digit possessing an extra phalanx. Sacral diapophyses cylinMaxillary teeth present. Thumbs opposable to other digits.
Reproduction aquatic; eggs are laid in a frothy mass. In Pseudis
paradoxa the tadpole may be nearly four times the length of adult.
drical.

FROGS OF COLOMBIA

— COCHRAN

AND GOIN

89

Genus Pseudis Wagler
1830.

Pseudis Wagler,

Generic diagnosis.

Fingers

free,

first

203 (type species, Rana paradoxa Linnaeus).

— Pupil

slightly free behind.

webbed;

p.

horizontal. Tongue circular, entire and
Vomerine teeth present. Tympanum distinct.

finger opposable to the others; toes completely

tips of all digits pointed or swollen into small disks.

metatarsals separated by a web.

Omosternum

cartilaginous;

Outer
sternum

a cartilaginous plate. Terminal phalanges simple.
Pseudis paradoxa nicefori,

Plates

new

subspecies

12g, 68a-c

—MLS 443, an adult from Monteria, Cdrdoba, Colombia,
22, 1966.
Rio San Jorge, between
Paratopes. — USNM 145076
Holotype.

collected

by Hno. Niceforo Maria, February
(larva),

Lagunas Puerco and Sabanas, Bolivar; USNM 145780 (larva),
Rioviejo, lower Rio Magdalena, Bolivar;
(larva), V&lledupar,
Badillo, upper Rio C6sar, Magdalena.
Diagnosis. Vomerine teeth in two large, transverse, well-separated
series between the choanae; greatest diameter of tympanum equal to
that of eye and equal to distance from eye to nostril; a thick glandular
ridge from posterior corner of eye, continuing above tympanum and

CNHM

shoulder to fade out behind the level of the axilla; lower surface of
femur with heavy gray reticulations enclosing pale round or oval

and sometimes irregular areas, with a wide pale stripe along anterior
surface of femur above the reticulations; lower surface of tibia pale

with a few thin gray vermiculations; venter olive-buff with some
more definitely aligned along center of
chest; a large triangular light spot outlined with gray below shoulder
attachment; belly immaculate; back with two very irregular longitudinal rows of darker spots on each side (one row near the midline,
the other in the dorsolateral area) a large light transverse spot
immediately behind anus and a second smaller patch behind it.
Description of holotype. Vomerine teeth in two very heavy,
transverse, well-separated series between the choanae; tongue twothirds as wide as mouth opening, deltoid, attached except on its
extreme border, which is straight; snout moderately short, rounded
when viewed from above, rounded in profile, the upper jaw extending
well beyond the lower. Nostrils superior, slightly projecting, their
distance from end of snout almost as great as their distance from eye.
Can thus rostralis indistinct; loreal region barely concave, sloping
broadly to the upper lip. Eye moderate in size, prominent, its diameter
two-thirds its distance from tip of snout; interorbital diameter twice
that of upper eyelid, equal to interval between nostrils. Tympanum
small gray spots on throat,

;

5 mm. venter olive-buff. Toes entirely webbed. pale olive-buff below. third toe longer than fifth. . metacarpal tubercles small but well developed. lower limb surfaces pale buff. fourth longer than either. no ventral disk. Fingers long. and a wide pale stripe above this reticulation along anterior surface of femur. Head and body. sides of body slate above. not hooked.5 15 mm. femur. a thick glandular ridge from head. slender. not enlarged. with a few small Color in alcohol. —Top tibia. heel reaches tip of snout. of pale dots invading the slate area. a thick tarsal ridge. chest.. foot. the first and second fingers with short lateral ridges. the webs pale but finely vermiculated with slate. no heavy skinfold across the chest. a heavy. 25 mm. one near midline. with fine transverse skinfolds across throat. enclosing round. with a heavy gray reticulation on lower femur. preceded by a longer transverse one reaching almost to middle of femur. knee and elbow overlap. a flat oval thumb pad present. separated from eye own diameter. a smaller round palmar callus.. a larger white patch below anus. head posterior corner of eye above — width.. first finger opposable to and shorter than second. free. palms of hands pale smoke-gray on inner two fingers. the other in dorsolateral area. belly immaculate olive-buff. toe disks slightly bulbous.. in postaxillary region equal to greatest When hind leg is adpressed. 27 mm.S. and belly. Body heavy skinfold on heel and width of stout. head length. when hind legs are bent at right angles to body. with some small gray spots on throat.. no outer one. heels touch. when limbs are laid along the sides. 17. its greatest by an interval equal to one -third its . the femur and tibia with indistinct darker bars. with a few thin gray vermiculations. 26 mm. head and back slate color with indistinct darker spots in irregular rows. 51 mm. ending near heel. more definitely aligned along center of chest. mm. a light irregular triangular spot outlined with dark gray below attachment of shoulder. soles of feet pale gray on three inner toes. Dimensions. venter smooth. posterior surface of femur slate-black with numerous small rounded white spots. reaching to base of penultimate phalanx of third tips of fingers large. slate color on outer two. oval and sometimes irregular pale areas. then merging with the slate color of loreal and subocular regions. but without granules except for a few small indistinct ones on sacrum. tympanum. 14. continuing above shoulder and fading out behind the level f axilla. hand. upper surfaces of limbs smoke-gray. whose tip reaches to center of penultimate phalanx of fourth.90 U. crescentic inner metatarsal tubercle.. dark on outer two. upper lip smoke-gray with a few faint darker spots. no external vocal sacs in the male. lower part of tibia and top of foot pale. a knee. NATIONAL MUSEUM BULLETIN 2 88 diameter equal to that of eye. Skin of upper parts heavily glandular.

more or less prominent. Number of vertebrae tending to be reduced. p. one-fourth webbed . to Colombian Species of Bufo Bony head ridges present. headcrests wide and swollen. Snout extended to a proboscis in adult. c1 . 1 . e2. d A subnasal ridge. Y/i times diameter of eye. B. No maxillary or vomerine teeth. 1 . The examination of the single adult from C6rdoba collected by Hno. B. and it gives us great pleasure to name it in for its collector. vulgaris Laurenti). Two large tadpoles from the Magdalena River drainage Bolivar have well-developed hind legs. headcrests including parietal strongly developed. — Generic diagnosis. with the original sacral vertebrae perhaps becoming incorporated into the urostyle.FROGS OF COLOMBIA —COCHRAN AND GOIN 91 — Remarks. or epicoracoids partly fused. parotoid oval. Family Bufonidae Shoulder girdle arciferal. rostratus nicefori Snout not proboscis-like. extension of postorbital ridge. rostratus rostratus Adpressed heel reaching axilla or front of shoulder. downward e1 . ceratophrys No b2. B. granulosus humboldti . The lower surface of the femur has two large. Sacral diapophyses dilated. ft 1 . . Toes moderately long. nostrils dorsal. one-half webbed Toes rather short. Bidder's organ present in male. A third tadpole. The receiving of the tadpoles from Bolivar (in 1961) and of the one from Magdalena was sufficient cause to postulate the occurrence of a new subspecies from northern Colombia. d Adpressed heel reaching angle of jaw or a little farther. Omosternum usually absent. mm c2 . horn on upper eyelid. A parotoid gland present. apparently resulting from a forward shift of the sacral articulation. toes about onefourth webbed. since the pattern of their lower femoral surfaces differed from the other forms of Pseudis paradoxa so well discussed by Gallardo (1961a). 1768. tympanum touching d2 . toes threefourths webbed. irregularly oval white spots in a very wide dark stripe. . 25 (type species. size to 35 mm . granulosus beebei . B. light-spotted dark stripe. projecting. size to 34 B. light. Genus Bufo Laurenti Bufo Laurenti. Key a 1 . No maxillary teeth. parotoid triangular. size to 42 mm. . not longer than eye. Upper eyelid with a hornlike projection. but often pointed and projecting in typhonitis. . has two wide irregular light subfemoral stripes invaded by a few dark spots and separated by a wide. . Lateral sacral processes dilated. Shoulder girdle arciferous. from the upper Rio Cesar in Magdalena. B. Niceforo Maria established the new form beyond a doubt.

sternosignatus a2 No bony ridges on top of head. 15. typhonius Angles of jaw not projecting. Skin granular to spinose. Description. moderate. haematiticus b2 First finger shorter than second. nostrils lateral or superolateral. mm . p. Eye large. angulate. size g to 43 B. a tarsal fold present. with than second and equal lateral ridges. mm . Bufo ceratophrys Boulenger. fig. and unnotched. ridges on top of head limited to rounded upper eyelids. /'. coniferus 2 Supraorbital and postorbital ridges forming a wide curve. p. not usually touching postorbital ridge.— Gorham. its diameter equal to its distance from tip of snout. b First finger as long as or longer than second. Bufo ceratophrys Boulenger Plate 14a-c 1882. size to 70 B. Fingers without disks. size to 23. . toes one-half webbed. soles of feet spinose. . marinus g 2 Skin glandular. c Tarsal fold lacking or indistinct. mm l . 179. d2 No .— Nieden. 22. first finger a trifle longer . snout 3-lobed when viewed from above (a central and one above each nostril). 50.— Gadow. their distance from end of snout one-fourth their distance from eye. blombergi g Parotoids moderate. loreal region flat. mm . g size to 90 B. Tongue its posterior border opening. Colombia. f2 Supraorbital and postorbital ridges forming a right angle. mm . size to 90 B. 1961. 141— Rivero. 2 (type locality. tarsometatarsal joint reaching beyond snout. Parotoids small. Nostrils lateral. tarsometatarsal joint reaching a point between eye and nostril. hypomelas el . size to 212 B. toes one-half webbed. 319. p. 96. Ecuador). .S. Can thus rostralis sharp. —MLS Caqueta. . p. 1901. 1 . Tympanum very small.5 mm. vertical. 1963. . slender. merging into the nearly vertical upper lip. size to 157 B. not very distinct. its greatest diameter apparently about one-third that of eye. or a little more. pi. . 22. Angles of jaw projecting in adult. truncate and slanting backwards in profile. p. the upper jaw extending well beyond the lower. 1 . separated from eye by an interval equal to free lobe. 1923. oval. B. mm . . very small but projecting. glaberrimus c2 Tarsal fold distinct. l .92 U. p. Bony crests at inner borders of one-third as wide as mouth a half-grown individual from Aserrio. size to 145 B. 1962. free. headcrests often greatly developed in old adults and involving the parotoids anteriorly. no tarsal fold. nearly its own diameter. NATIONAL MUSEUM BULLETIN 2 88 tympanum subnasal ridge.— Gallardo. 1882a. prominent. toes one-fourth to one-third webbed. nearly smooth. head ridges heavy but not unusually e2 developed. . mm . Parotoids large. interorbital diameter about three-fourths that of upper eyelid (including the greatly prolonged "horn" on outer edge of eyelid) and equal to interval between nostrils.

widening to an elongate parotoid gland.— —COCHRAN FROGS OF COLOMBIA AND GOIN 93 which reaches to base of penultimate phalanx of third. soles granular.12.. Bufo rostratus Noble. — Gorham. hand.12. heels touch.5 mm..5 mm.5 — Head and body. Dimensions. similar. Skin of upper parts with minute tubercles and granules. 23 mm. femur. no external vocal to fourth. although in the type it is somewhat more faded than in the fresher Colombian specimen. knee and elbow slightly overlap. on top of head.5. 8 foot. and becoming narrower but still quite distinct and set with tubercles from axilla to groin. when hind legs are bent at right angles to body. no ventral disk. heel reaches posterior corner of eye. Santa Rita Creek. EDUADOR: BM MLS 50.. belly and lower proximal femur distinctly but finely granular. a slight skinfold across the chest. p.) but otherwise the two specimens are very is strikingly similar in the two specimens. north of Mesopotamia. 14 mi. 1962.. Bufo rostratus rostratus Noble Plate 14d-f 1920. — Upon direct comparison. 80. head mm. third toe a trifle longer than fifth. the dorsal pattern Specimens Examined COLOMBIA Caqueta: Asserfo. a pronounced skinfold on heel and knee. Colombia). but this also probably is due to age. its place being taken by the parotoid and lateral tubercles.. Antioquia. 10. p. a small. 16. a heavy lateral fold beginning at posterior corner of eye. ending at heel. throat and chest faintly granular. pointed metatarsal tubercle and a smaller round outer one. but this most likely is due to its greater age. 96. and on sacrum. and a similar but smaller inner one. The bandings on the arms and legs of the two specimens are essentially identical. enlarged along middorsal line. Gallardo. in postaxillary region equal to greatest width of head. head mm.151) are certainly conspecific.151 (type). width. 8. a distinct outer tarsal ridge edged with small tubercles. no glandular ridge from posterior corner of eye above tympanum. . Remarks. tibia. p. its tip reaching to base of antepenultimate phalanx of fourth. webbed at the base. 445 (type locality. The parotoid glands on the type are more conspicuous. Toes without disks. 80. 8 mm. a small rounded palmar callus. Even softer. metacarpal tubercles well developed. 10 mm. When hind leg is adpressed. when limbs are laid along the sides. Body short and stout. an oval projecting thumb pad present. 6. sacs.. and now somewhat MLS length. 50 and the type The type is larger (35 (BM mm. 1963. The bony ridge on the inner and posterior sides of the orbit is more pronounced in the type.5.

interorbital diameter 2% times that of upper eyelid almost twice the interval between nostrils.. first finger shorter than second. Nostrils lateral. But Noble's parotoid glands . Fingers one. dividing at posterior inner edge of eyelid. when limbs are laid along the sides. 13 mm. its posterior border free and slightly notched. femur. projecting. Eye large prominent its diameter two-third its distance from tip of snout. When hind leg is adpressed. Canthus rostralis very sharp.5 mm. foot. Tympanum hidden. a round palmar callus. and on upper limb surfaces. tongue very long and slender. Description. the upper jaw extending far beyond the lower due to the elongate snout tip. Dimensions. 36. soles granular. venter finely granular everywhere. their distance from end of snout equal to their distance from eye. the latter more apparent between shoulders. 14 miles north of Mesopotamia.... the proximal ones divided. the inner ridges converging and disappearing on the back above the shoulders. Head and body. turning backwards at a right angle above tympanum. Colombia. 9 mm. their tips reaching to base of antepenultimate phalanx of 4th toe.5 mm. 13 mm. Color in alcohol (from Noble's original description of the type) Color above yellowish brown. female paratype from Santa Rita Creek. Body stout. Toes short. 3rd and 5th toes subequal. heel reaches to angle of jaw.— 94 U. when hind legs are bent at right angles to body. knee and elbow are separated. and continuing as a series of sharp. no skinfold across chest. hand. snout pointed when viewed from above and in profile. ventral surfaces yellowish white reticulated with dark brown. Skin of upper parts with small granules and scattered larger conical tubercles. diverging to border inside of upper eyelid. less than half as wide as mouth opening. with lateral ridges.. . Heavy bony ridges on top of head beginning at prolonged tip of snout. an oval thumb pad present. heels are slightly separated. sloping a little to meet the upper lip. an external vocal sac in the male. no ventral disk.S. on sacrum. vertical. ovate. an oval inner metatarsal tubercle. in postaxillary region a little less than greatest width of head. and reaching nearly to penultimate phalanx of third. head length. three-fourth webbed. an outer tarsal ridge bordered with conical tubercles. 10 mm. loreal region concave.5 mm. indistinctly marked with darker brown. In Noble's diagnosis. forming crossbands on the legs. NATIONAL MUSEUM BULLETIN 2 88 —AMNH 1384. metacarpal tubercles well developed. a series of skinfolds on heel and knee. head width. 13. Antioquia.. and an outer one nearly equal to it in size. — — rather small. the outer ridges bordering the eye behind transversely. sub triangular. 13.third webbed. tibia. the type is said to have the fingers one-third webbed and the toes three-fourths webbed. Remarks. conical tubercles along side of body to groin. fourth longer than either.

— Description of holotype. lower surface of proboscis smooth and rounded. Bufo rostratus nicefori. an adult from El Chaquiro." This "narrow seam" has been called the "lateral ridge" in the present paper. . the upper jaw extending well beyond the lower. — USNM 163476. nearly parallel crests beginning between the eyes and continuing back on the parietal region. its diameter slightly less than its distance from tip of snout. the tip elongate and rounded when viewed from above. tongue about half as wide as mouth opening. its posterior border free and unnotched. Canthus rostralis swollen. A pair of heavy. prominent. interorbital diameter twice that of upper eyelid. Specimens Examined COLOMBIA Antioqtjia: Santa Rita Creek. Hind leg short. we shall have a broader conception of the . When Myers and Leviton publish the results of their current studies on coniferus and its allies. Eye large. . their distance from end of snout equal to their distance from eye. Noble further states that the toes are "about three-fourths webbed. new subspecies Plate 15d-p Holotype. AMNH 1359 (holotype). Nostrils lateral. distinctly webbed at base. 1384 (paratype).FROGS OF COLOMBIA —COCHRAN AND GOIN 95 detailed description states that the fingers are "webbed only a little beyond the metacarpals. loreal region weakly concave. north of Mesopotamia." as a seam to the tips of all the toes Noble believed that his rostratus was related to Bufo coniferus Cope. Dorsolateral tubercles large and blunt. bluntly pointed in profile. Paratypes. heavy. same data as holotype. 14 mi. greater than interval between nostrils. the web extending to the end of the phalanx of the two inner toes. Diagnosis. Toes one-fourth to one-third webbed. Finger short. adpressed heel reaching axilla or front of shoulder. supraorbital ridges coarse and heavy. Tympanum hidden. NicSforo Maria in December 1966. status of these toads. slightly projecting. Tympanum covered. extending to groin. very elongate. Fingers fairly short . Antioquia. the web continued along the edges of the fingers as a narrow seam. top of head between them appearing collected concave. MLS 449-52. snout proboscis-like. continuous with the rostrum. and to the base of the antepenultimate of the fourth toe. but the web extending into the metacarpal region making the fingers appear about one-third webbed. — — by Hno. web continued . Snout bluntly pointed at tip and projecting. to the base of the last phalanx of the third and fifth toes. sloping a little towards the upper lip.

two less-distinct rows are on each side of midline) venter granular. a very thick glandular ridge from posterior corner of eye ending in the short round parotoid. —Dorsum immaculate clove brown. a small rounded present. femur.. with faint traces of a web. [The collector notes that the two toads were nearly black when alive.. in the rostratus forms it is the nasal even an unusually long-snouted typhonius can scarcely be mistaken for a form of rostratus. width. Body stout. third and fifth toes subequal. Numerous differences exist. foot. The back is very dark in all the individuals. The belly may be mottled seal brown and sepia.. 34 mm. and. heels are separated.. reaching halfway on penultimate phalanx of third.. no skinfold across the chest. first finger shorter than second. .. side body clove brown to sepia. a larger flat palmar callus. a small round inner metatarsal tubercle and a smaller outer one. tibia. head length. and three have narrow light mid dorsal lines. one-fourth webbed. no ventral disk. knee and elbow are separated. howin old adults. Dimensions. When hind leg is adpressed. in postaxillary region equal to greatest width of head. The four paratypes are much like the type in structure. a thick skinfold on heel and knee.5 10 mm. Bujo ceratophrys Andersson and B. no apparent tarsal ridge. 8. with large and small granules mixed on the belly and very small ones on throat. palms of hands and soles of feet dull sepia. flattened but without true lateral ridges. In typhonius it is the post- ocular and supratympanic crests that are frequently over-developed and supraocular regions that show most specialization.] Remarks. with none of the dorsal — spotting characteristic of Bujo typhonius. typhonius and B. Skin of thumb pad upper parts thickly covered with rounded tubercles in three rows (the largest tubercles in a row along dorsolateral line. hand. or olive to dull buff with somewhat more distinct dark dots. Color in alcohol.5 mm. external vocal sac in the male.5 — Head and body. their tips reaching to base of antepenultimate phalanx of fourth. upon close examination. 11 mm. posterior femur clove brown above lightening to drab below. throat paler in the center.96 NATIONAL MUSEUM BULLETIN 288 U. belly wood brown with of head and sepia suffusions. short. dapsilis Myers and Carvalho likewise suggest an interrelationship with B. when hind legs are bent at right angles to body. mm. metacarpal tubercles not very distinct. heel reaches axilla. a median . darkening to clove brown around edge of jaw. when limbs are laid along the sides. rostratus.S. 11. One of the paratypes appears to have a partial light midventral line. 11. 12 mm. broad. ever. head mm. Toes broad. except that in one example the adpressed heel reaches the front of the shoulder and the toes are one-third webbed. fourth longer than either.

— Lutz. Trinidad. 1952. Bony — MZUM 2.— Beebe. p. rounded in profile. descending nearly vertically to the upper lip. I. one-fourth webbed. its tip reaching to base of antepenultimate phalanx of third. Canthus rostralis concave. figs. interorbital diameter 1% times that of upper eyelid and twice the interval between nostrils. Tongue two-fifths as wide as mouth opening. pi. rostralus nicefori are extremely short. 1933. its disk reaching to center of antepenultimate phalanx of fourth. free. metacarpal tubercles well developed. separated from eye by an interval equal to half its own diameter. snout broadly rounded at the tip when viewed from above. venter finely granular. Nostrils more dorsal than lateral. 5. globulosus (not of Spix). third toe longer than fifth. oval.). a skinfold on heel and knee. from Gaira. 124 (part). B. 38. and (a short one) on parietal region. 1927. 1936. 6. — Ruthven. Body stout. 1959. knee and elbow are widely separated. 1927.— Lynn. heel reaches axilla. Eye large. Tympanum distinct. W. Skin of upper parts granular. Description. 113. its greatest diameter one-half that of eye. and a smaller oval outer one. 174. although the proportions of its femur and tibia to total length are overlapped by the Ecuadorean B. Fingers moderately long. Bufo granulosus beebei Gallardo. Magdalena. the distal ones mostly double. Toes rather short. 25. 1. no distinct tarsal ridge. Churchill-Roosevelt Highway. 16. a small oval thumb pad present. pi. . first finger shorter than second. the upper jaw extending far beyond the lower. When hind leg is adpressed. ridges on top of head encircling eye. Bufo granulosus beebei Gallardo Plate 68d-f 1922. 1927. 1. loreal region concave. 6. figs.— Parker. its diameter equal to its distance from tip of snout. 11. p. 1934. along canthus and nostril. a much larger round palmar callus. 45527. pi. their distance from end of snout about one-half their distance from eye. projecting. p. p. 1922. p. with weak lateral ridges. Bufo 1965. 114. p. heels touch. — Lutz. Nominate rostratus has longer legs. p. in postaxillary region equal to greatest width of head. Bufo granulosus (not of Spix). 8. 5. figs. a small pointed oval inner metatarsal tubercle. the granules on head and along dorsolateral region larger. p. fourth longer than either. p. below above and below tympanum. 42. dapsilis. when limbs are laid along the sides. Colombia.FROGS OF COLOMBIA The —COCHRAN AND GOIN 97 legs of B. its posterior border free and unnotched. with a large patch of fine irregular wrinkles on posterior part of belly and on proximal femoral surface. prominent. maps (type locality. when hind legs are bent at right angles to body. 55 (part). with femoral and tibial lengths falling below the extreme lower limits of the others.

mm. head height. 55 (part) . Head and body. upper limb surfaces drab with sepia markings on top of femur. the spicules around eye. Cope. — locality. hand..—Noble. p.S.5 mm. no ventral disk. an external median vocal sac in the male. foot. Tolima. USNM 146815. and numerous smaller sepia spots surrounding the enlarged tubercles along dorsolateral region and nape. Description. head . 9. a few pointed spicules below tympanum and beneath eye. VENEZUELA: Maracay.. the tips and tubercles darker. Bufo granulosus humboldti Gallardo Plate 13a-c — 1899.. west of Girardot. p. 1899. is really of about the same proportion in both forms in relation to percentage of head-and-body length. 1924. p. while the head height tends to be a little more than in beebei although they slightly overlap. Dimensions. ending at parotoid.5 — Dorsum wood brown with a pair of small oval Color in alcohol. beebei are partially confirmed by comparing critical measurements of head width and head height. with a prolongation forward and inward along the canthus rostralis. 14.— Ruthven. In humboldti the head width is a little greater. sepia spots between the shoulders. 43 width. tibia. 14 mm. and . an adult male from Villavicencio. and upper lip dark. 1. 6. 1922. posterior surface of femur drab to clay color with two or three indistinct darker spots. 66. —Some of the minute differences by which Gallardo (1965) distinguishes B. g. maps (type Gualanday. Colombia) Hendrickson. Bufo granulosus (not of Spix). Augustine. USNM 141533. and two large sepia spots on upper surface of tibia and foot. said to be "elongate" in beebei and "short" in humboldti.— Dunn. 513.— Stebbins and 1965. no skinfold across the chest. — mm. p. A narrow but sharp bony ridge encircling eye. nostril. 2 88 a heavy glandular ridge from posterior corner of eye above tympanum. TRINIDAD: St.. MZUM 45527. 1944. another crescent-shaped pair at the beginning of the sacrum. Remarks. p. head . femur. p. which is small and nearly triangular. 117. 11 mm. Bufo granulosus humboldti Gallardo. length. Specimens Examined COLOMBIA Magdalen a: Gaira. fig..5 mm. 514. a fine ridge along rim of upper jaw.. side of body drab above fading to dull clay color below with a few indistinct sepia spots in upper part. Meta. 1959. 15. 12 mm. 98 NATIONAL MUSEUM BULLETIN U. 14 mm. side of head ecru drab. Colombia. USNM 97193-5. The snout. granulosus humboldti from B. palms of hands and soles of feet pale clay color. venter immaculate clay color.

its greatest diameter one-half that of eye. when hind legs are bent at right angles to body. Dimensions. hand.. metacarpal tubercles well developed.5 mm. skinfold slightly longer than eye. with transverse folds across abdomen. when limbs are laid along the sides..5 mm.. Toes moderately long. with lateral ridges. side of (in this specimen) except for a dark suffusion on the throat of the male. their distance from end of snout one-half their distance from eye. heel reaches just beyond axilla. loreal region concave. its diameter 1% times its distance from tip of snout. femur. its posterior border free and unnotched. ending at parotoid. immaculate are chevron-shaped on the back. When hind leg is adpressed. one-half webbed. 10. but touching postorbital crest. snout truncate at tip when viewed from above. another narrow crest bordering upper lip from eye to snout tip.. the upper jaw extending well beyond the lower. the crests body wood brown with small dark brown spots fading out below. no skinfold across chest. Eye large. first fingers of male are roughened in breeding season. parotoid gland short. 14. Snout short. broadly triangular. a short coarse glandular ridge from posterior corner of eye above tympanum. reaching to base of penultimate phalanx of third. its disk reaching nearly to base of antepenultimate phalanx of fourth. heels overlap. separated from eye by an interval equal to about one-third its own diameter. first and second fingers subequal. 41 mm. projecting in profile. Head and body. fourth finger slightly longer than either. Nostrils dorsal. in postaxillary region equal to greatest width of head. Fingers fairly long. venter finely granular. much greater than interval between nostrils. head — width. knee and elbow are slightly separated. tongue about one-third as wide a mouth opening. head length. the tubercles on top of head smaller and flatter. encircling eye. wood brown with foot. Tympanum distinct.5 —Dorsum brown spots which are more mm. 14. limbs pale wood brown above with coarse clove brown spots.5 mm. Body stout. slanting outwards to the upper lip. a small oval inner metatarsal tubercle and a smaller round outer one. strongly projecting.. tibia. rounded. Can thus rostralis sharp. third toe slightly longer than fifth. with a subnasal ridge. Skin of upper parts tubercular. oval. which . curved. no distinct tarsal ridge. no ventral disk. 13 mm. a small oval thumb pad present. Color in alcohol. a heavy on heel and knee. soles granular. continuing as a broad but weak posterior occipital crest. side of head outlined in clove brown.. a large round palmar callus. venter light wood brown. prominent. 13. a median external vocal sac in the male. large irregular clove or less transverse between the eyes and wood brown. a narrow crest curving inward from nostrils then outward to anterior eye. 15 mm. faintly webbed at base. interorbital diameter 1% times that of upper eyelid.FROGS OF COLOMBIA — COCHRAN AND GOIN 99 another back of eye to the small parotoid.

146201. Cailo Guapaya. Boca del Rio Guaviare. Specimens Examined COLOMBIA Antioquia: Casabe. USNM 144153. The males can be distinguished by the brown patches of rough scales on the inner surface of the first finger in breeding season. Differences are very slight between these two forms. In some males the vocal sac extends as far back as the armpits. soles of feet and palms of hands ecru drab. 150151-3. The peculiar shape of the snout in granulosus is likewise an identifying character. 146816-7. 100 m.S. Magdalena. USNM 150733-41. tibia. Canal Zone. USNM 148135. Fort Kobbe. northeast of Villavicencio.. the toe tips and tubercles sepia. USNM 144574-83. USNM 117515. the smallest specimen to display the brown patches is USNM 146636.5 millimeters in total length. foot. These forms have been studied most recently by Gallardo. 7 km. The larger and more circular tympanum also is a prevalent characteristic of the male of this subspecies. 146635-6. BoLfvAR: Sincelejo to Tolu. USNM 140493. USNM 53739. lower Rio Manzanares. Only the comparison of large series of beebei with humboldti can confirm — or repudiate their validity. Sierra de La Macarena. The very short femur.. Vaupes: Arnanaven. Cristalina. Finca El Aranar. between Santa Marta and Mamatoca. Sierra Macuira. USNM 144573. USNM 151292-302. Sabana. 380 m. measuring 38. who has called most of the Colombian toads humboldti. Tocaima. The species has several distinct forms in different regions of South America." Villavicencio. and hand distinguish members of the granulosus group from other Colombian toads. central. Valle: Rio Calima near C6rdoba. and forearm. One-half mile south of Pajaro. USNM USNM Norte USNM Tolima: Mariquita. edge of Cano Losada. PANAMA: Rio Calobre. USNM USNM 144571. USNM 150892. and southern states of Colombia. 146815. C6rdoba: Cano Betancf. Remarks. USNM Guajira: USNM Junquilla. 30-60 m. de Santander: Astillero. 147266-7. with the single exception of one from Guaira. "La USNM 144572. but the former subspecies has a slightly greater head width and head height.100 U. Cundinamarca: Melgar. . which he records as beebei. USNM 144584. USNM 145773. Atlantico: Rio C&ar. Piojo. Melgar. Bonda. USNM 146814. The dark spotting of the belly apparently has no regional significance. posterior femur wood brown with dark spots which fade out below. as both spotted and unspotted bellies occur in toads from the northern. USNM Meta: 152735-49. In the present series. NATIONAL MUSEUM BULLETIN 288 form three irregular crossbars on tibia. El Cucharo. 146385-9. f USNM Magdalena: Cienaga. 115380. 152688-93.

Brazil). 1882a.— Daudm. 19. [61]. p. p.—PersiCca. fig. 181. 292. figs. p. —Schneider. 299. pp. p. p. 3. 1843. p. 1873a. 1925. margaritifer. Schmidt. 1939. 15. Brazil). p.— Melin. p.— Beebe.— Stebbins and Hendrickson. 61— Mello-Leitao. 425. 1838. 1858.— Parker. 47. 11. — Miranda. 153. 1921. p. p. fig. 1911. 178. p.— Gorham. 14. 143.— Dumeril and Bibron. 1898a. 1862b. 1871. 159. 1824. 554.— Muller and Hellmich. p. p. Amazon River). 2 (type locality.— Boulenger. p. Trachycara fusca Tschudi. 253. 509. 1896. Bufo margaritifer. p. p. 32. p. Peters. 67. 1159.— Merrem. 20. 1912. p. 50.— Procter.Riberiro. 1923. northeast Peru).— De Witte. America). 439. p. p. Otolophus — Fitzinger. 1875.— Fowler. fig. 1913.— Dunn. p. p.— Lidth. 20. 1050. 156. 317. 14. fig. 69. p. 1799. 1926. 1874b. p. 1927. 1913. — Gmelin. 29.— Cope. Lutz. 1959.— Gans. Bufo thyponius. 207. 1050. Bufo typhonius. p. 1871. p. 635. 1824. pi. 1843. 125. p.— Ahl. p. Brazil). 142.. 1802. pi. . 1919. p. Rio Solomoes. p. 33. 135.— Steindachner. Spix. 89. 52. 5 (type locality. 600. p.— Hellmich. 1926. 171. typhonius. 1824. 4 (type locality. — Tschudi. 1838. p. 1904. 108. p. Eurhina proboscideus. 1914. p. 1022. 1858. 1963.— Peracca. p. p. p. p. 1824. p.— Carvalho. 1858. — — 1824. 139.— Goin and Layne. 1939a. 21. p. p. 211 (type locality. fig.— Werner. p. 1875. 17-19 (type locality. Steindachner. 21.— Ruthven. p. 1858. 1944a. 43. 1887a. 1896. p.— Despax. p.— A. p. 1958. 104. p.— Andersson. Amazon River). 1931. 132. figs. fig. p. fig. p. Peru). 252. 80. 1824. 96. Brazil). 1929. 217 (type locality. 1901. 2. p. Oxyrhynchus acutirostris Spix.— Nieden. 16. 1952. pi. 78. 16. 1929. 1-lb. p. 93. pi. 1945.— Cott.— Travassos and Freitas. photograph. Bolivia to Peru). 30 (type locality. 1802. Otilophus 1845. Bufo margaritiferus. p. p. 624. p. 1962. 266. p. p. 1944c. p. p. Espada. p. p.— Baumann.— Lutz and Kloss. p. 123. 1867. Oxyrhynchus nasutus. 1873e. 357. region of Ceja between Jauja and Uchubamba. 1799. 1937. 6. p. p. 537. p. 1930b. Oxyrhynchus naricus Spix. p. p. 1930. p. Gunther. Oxyrhynchus spixii Wied 1824a. 637. 718.— Brazil and Vellard. 40.— FROGS OF COLOMBIA —COCHRAN AND GOIN 101 Bufo typhonius (Linnaeus) Plate 14g-i 1768. 280. 1789. p. 1858. p. 89. 1841. 49. fig. 282.— Hno. pi. 1960. of Tarma. p. 12. 132. 1920. 403. pi. Nice7oro-Marfa. — Espada. 1875. 69. 17. 1820.— Gunther. 52. 1. 1758. p. 1843. 36. 88. p. Rana margaritifera Laurenti. 1904. p. fig. p. Schmidt. 102. 1799. pi.— Shaw.— Crawford. 16. Bufo pleuropterus O. 1789. 179. 1802. 1901. p. 1933. p. 118.— Gunther. 519. A. pp. 251. 1843. — — 1875. 1901. 1884a. fig. 1903a. Oxyrhynchus typhonius.— Barbour and Noble. 226. p. 32. 2. O. 191. p. Oxyrhynchus proboscideus Spix. p. p.— Gines.— Gallardo. 185 (type locality. 97. — Gmelin. 1919. 3 (type locality. — Osilophus typhonius. 513. — pi. Bufo nasutus Schneider. 1942. p. 1941. pi. 8. 1935. 1939. 1959. 1870. 1944d. pi. p.— Crawford and Jones. 1898b. fig. 120. pi. 1938.. p. figs. p. 337-262—70 8 p. Cope. 45. p. 207. 1936. p. 1926. — Fitzinger. 130. Rana typhonia Linnaeus. Cope. — —Peters. 1934. 1870. Otilophus margaritifer. 1917. p. 111. 1926. 3 (type locality. 156. Oxyrhynchus iserni Espada. 69. Peters. 1934c. 1859a. 399. p. 23. 1803. 45. 149.

from Puerto Narifio. Cei. Amazonas. heel reaches center of tympanum. third adpressed. posterior part of parotoid not involved in the enlarged crest. knee and elbow are slightly separated. when hind bent at right angles to body. Skin of upper parts covered with fine and coarse granules. Bufo typhonius typhonius. 1953. Toes moderate. heels are widely separated. its diameter nearly as great as its distance from tip of snout. 29 p.102 1933. and lower limb surfaces. snout pointed when viewed from above. and ending above and behind the axilla. p. elongate oval. 8. Fingers long and slender. first finger a little longer than second. legs are . the upper jaw extending well beyond the lower. both reaching nearly to base of antepenultimate phalanx of fourth toe. Nostrils opening upwards. When hind leg is lateral ridges. Bufo (Otilophus) typhonius. its upper rim involved in the greatly enlarged crest. venter finely granular on chin. p. Body stout. Bony ridges on top of head greatly enlarged beginning with supraorbital crest and extending continuously as postorbital crests behind the eye and above the ear. soles spinose. fourth longer than either. slightly projecting. no inner tarsal ridge. metacarpal tubercles well developed. NATIONAL MUSEUM BULLETIN 2 88 — Leavitt. its greatest diameter three-fifths that of eye. their distance from end of snout one-half their distance from eye. its posterior border free and unnotched. a weak parietal ridge bifurcating from the posterior part of the supraorbital crest. all the digits thumb pad with bulbous. an internal vocal sac in the male. no ventral disk. much greater than interval between nostrils. a flattened oval inner metatarsal tubercle and a smaller roimded outer one. separated from eye by an interval equal to one-half its own diameter. Tympanum distinct. no skinfold across the chest. belly. but merging with a longitudinal line of conical glands which are continued along the side to the groin. Colombia. — Description An adult male. — Rivero. descending nearly vertically to the upper lip. but a weak outer one to heel. U. nearly one-half webbed and with and fifth subequal. Can thus rostralis sharply ridged. interorbital diameter more than twice that of upper eyelid. in postaxillary region equal to width of head without the knob-like protuberances on jaw. reaching their greatest height just before merging with the large parotoid. 1933. prominent. USNM 146824.S. 511. more coarsely granular on chest. with lateral ridges and weak basal webs. swollen tips. a larger oval palmar callus. a knoblike protuberance on angle of jaw. truncate in profile. becoming much larger posteriorly. loreal region flat. — 1953. a middorsal series of seven large button-like prolongations of the lumbar vertebrae (in old males only). 1961. reaching nearly to base of penultimate phalanx of third. a distinct oval present. Tongue nearly one-half as wide as mouth opening. a slight skinfold on heel and knee. Eye large. when limbs are laid along the sides.

In fact. while the inner part is a contrasting pale olive-buff. palms of hands light drab. side of head light drab. aside from the hyperdevelopment of the head crests. USNM 146827 (46. femur. the tubercles and disks fawn color. 42). Dorsum vandyke to seal brown. Amazonas. the tubercles lighter. only slightly so and quite distinctly so in the remaining three specimens of the series.. One of the most peculiar characteristics of this toad.FROGS OF COLOMBIA — COCHRAN AND COIN 103 — Dimensions. the knob is very distinct in more than three-fourths of the toads at hand from various regions in Colombia. the knob is well developed in 26 of 30 examples from Amazonas and 14 of 16 examples in three of the others. One of the old males (USNM 146823) has two black lunar markings between the eyes and three other pairs of such markings on each side of the midline of the — — back. The males at breeding time have the area on the inside of the first finger roughened. side of body drab below. The toes are one-third to one-half webbed in all Colombian toads examined.5 mm. externally. as in USNM USNM 146830. indistinct clove brown spots on back and slightly darker edges on the crests. The smaller toads from Puerto Narino. Head and body.. venter light drab-gray with some large irregular drab spots on the belly. 20. while the "buttons" along the backbone are evident in four of the specimens in the series. figured by Cochran. the webs and disks a little lighter. and forearm. hand. tibia. 68 mm. p. head length. has five large and two small "buttons" and otherwise is similar to the one from Colombia described above. The belly may be heavily brown-spotted.. This characteristic occurs in its most extreme form in old males with large head crests. soles of feet vandyke brown. The knob at the angle of the jaw is produced to varying degrees in all. they can be distinguished. 1961a. 26 mm.5 mm. is the appearance of a row of five to seven "buttons" on the midline of the back coinciding with the upper central part of the seven vertebrae.. tibia. 26 mm. from Meta. 26 mm. foot. as in or nearly uniform olive-buff. 22. posterior surface of femur hair brown. Remarks. from the females even in a melanistic population.. An example from the Maroni River. British Guiana (USNM 108987.5 mm. The lower surface of the tarsus and outer part of the metatarsal region are vandyke brown. and very pale suffusions of light drab on chest and throat. are more brightly marked.. foot. 16 mm. head width. immaculate. except for a few Color in alcohol. in length) having a wide pale middorsal stripe and showing two wide dark crossbands across femur. merging gradually with the darker brown of the upper surfaces. The protrusion of ribs through the skin is known in a few . For instance. 146828.

Macarena. Choc6: of Rfo Andagoya.. "Darien. Cauca: Near Moscopan. Villavicencio. USNM 144664-6. above Puerto Asfs on the upper Guamues. Urcusique. Meta: Buenavista." USNM Cundinamarca: Beltran. 144626146823-30. USNM 147399-400. Antioquia: Medellin. Guaicaramo. roqueanus Melin. Camp No. ci6n. Puerto Narifio. such as the Spanish newt. San Francisco. Camp "Las Ranas. 81343-8. over its extremely wide range. 1913. USNM 14624-31. Arteaga BolIvar: Cienaga de la Raya. 400 m. is extremely have — variable in the bony development of its head crests and vertebral knobs. Putumayo: San Antonio. is not considered here.— 104 NATIONAL MUSEUM BULLETIN 288 U. tributary of Rfo Baud6. Rfo San Juan.500 m. USNM 146821-2 (female with eggs) La Macarena. 145776. USNM 144661-3. Sinu River side.. Rio Mataja. distinctly "knobby" vertebrae to name only two it is perhaps not surprising that an overgrowth on the back also should take place in a species where abnormal growth is already evident in the postorbital crests. 61777. 146200. 500 m. Camp "La Macarena.) 150154-6. 330 m. 144645-6. Villa Maria. and B. tributary of Rfo Opogodo. 1941. As some toads Bujo alvarius and B. La Mesa. upper Rio Apaporis. valliceps. 61680. Villa 63841. it is another of the forms requiring comparison of many specimens from Central and South America before a discussion of some proposed subspecies becomes practicable. 147142-7. tributary 61696of the upper Putumayo.S. upper Rfo Giiejar. 61721-30. Regenera61695. 61691-4. chanchanensis Fowler. USNM Pep<§. Rfo Guapaya." USNM 144658. For this reason the status of Bujo alatus Thominot. 150623-5. Leticia. Uraba. CNHM CNHM 144659-60." USNM 144657. C6hdoba: Alto de Quimarf. Amazonas: USNM USNM USNM CNHM CNHM CNHM USNM CNHM CNHM USNM USNM 3 (juv. Pleurodeles waltl. La Guayacana. 1. Serranfa de La Macarena. Cano Sando. 61757-63. UF Gino-Goj6. 146812Caqueta: Aserrio. 33. 61766. 1. 10017.." lower Rfo Baud6. Since this species. USNM 147276-316. -8(?). 150157-63. 144931-2 (juv. near Quebrada Santa Anna. Las Mesetas. Rfo Mira. 63842. amphibians. upper Rfo Magdalena. USNM 145107- 23. USNM 144653-5. Murucucu. 350 m.. Puri. 10-15 mi. USNM USNM USNM USNM CNHM and Puerto Boyaca.. Rio Apaporis. USNM 144656. probably upper Rio Sinu. Sorotama. USNM 144650. Umbrfa. between Girardot . USNM 147253. Camp "Las Ranas. USNM 144651-2. lower Rfo Apaporis. 144634-6. USNM 144638-42. MLS 51. USNM 144647-9. 1884. USNM 144667. Rfo Orteguaza. Rfo Renjifo. 250 m. tributary 4337144643-4. upper Rfo Pepe.. 124266. 248. USNM 144673. Gorgona Island. 81339-42. B. Fusagasuga. 127856-62.). Nari5jo: Imbilf CNHM CNHM . west of Playa de Oro. Specimens Examined COLOMBIA USNM 144621-5. mouth of Cano Losada. 8. 63843-4.

1930. p. 1882a. — 195. 1 mi. 144668-72. Bufo marinus. 1959. p.— Spix. lazarus.6 USNM 142107-9. lower Calima River. p. 65. p. Docidophryne — Fitzinger. — Laurenti. Gir6n. 1946. 15 Wied.— —COCHRAN AND GOIN FROGS OF COLOMBIA 105 USNM Lebrfja. La Uni6n. 1948. 1961. Quebrados Banos. Valle: El Pailon. 145082. p. 1799. — . p. Bufo brasiliensis Laurenti. Amazonfluss). 1963. 1953.— Boulenger. 96. Bufo marinus marinus. 1952a. Rio Raposo. p. 1862. 1920.— Baumann. 1867. pi. 80. p. p. 315.— Cei.] Colombia. p.— Berg.. 151332-77. p. 1768. 211 (type locality. Merrem. 102. 1858. 1860. Bufo horridus Daudin. 1953. 1921.— Gallardo. Bufo pythecodactylus. 46. p.— Nieden. 358. Bufo marinus agua. 1799. from Buenaventura. 511. above San Miguel.— Girard. 415. p. 1843. Beni. 425. p. 26. p. Goin and Layne. 1858. 1768. near Campo Santa Clara. 1952. p. Bombinator horridus. America). p. Crawford. 513. 1899. p. — — — 1802. 15. p. p. 14. [Cundinamarca. p. — — Rivero. 1802. 1862b. pi. 209.— Steindachner. p. p. Campo Santa Clara. 97. Rana marina Linnaeus. Vellard. p. p. p. 27. 1958. 1920. east of Cocota. p. p. 703. 1 (type Docidophryne agua. p. 1820. USNM 107648. p. Virology Field Station. 27. 1901. p. . Vanzolini.— Giinther. pi. — 12 (type Bufo molitor Tschudi. 5. Bufo humeralis Daudin. pi. 148. pi. 160. fig. 25. 219. 38. 1961.— Dunn. p. 43. Cayenne). "l'Amerique meridionale principalement Br&il") 1803. USNM 127146. 17. 6. 36 (type locality unknown). 1896. 1912. 144674-7. 609. p. 1961. 1845. fig. pi. Bufo marinis. 226. 7. pi. 7935-6. 1-3. Camp Carton de Colombia. Barbour. Santander: CM USNM USNM USNM USNM USNM USNM USNM USNM 123963. p. 1875. Pucallpa. km. 1959. 201. 1824.— Ginfe. p. Cercado. p. Bufo lazarus Spix. figs. — Cope. 224 Johnson. p. 1824. p. 178. p.— Freiberg. pi. 1843. 1820. 250. San Gil. — Shaw. 147517-8. (type locality not given). p. p. Solimoens. 96. p. 1901. 1931. 190. 179. 394. p. USNM 127144-5. 1952. 37 (type locality.— Breder. Vdlez. 26 p. 1841. Barbour and Noble. locality. 205 (type locality. p. 44. Anchicaya. 73. 1901. 32.— Dumeril and Bibron. Bufo maculiventris Spix. 1962. Bufo pithecodaclylus Werner. 45. Peters. 1860. Fitzinger. Santa Cruz. 168. Rfo 144678. Quebrada La Lechera. 1941. 114. Phrynoidis agua.— Schneider. 1956. p. — Brazil). p. — Merrem. Gadow. Bufo marinus (Linnaeus) Plate 13p-h 1758. 1923. 1803. 1944c. Tolima: Mariquita.—Jimmez de la Espada. 31. figs. 138 Lutz and Lutz. 118— Gorham. p. 1 (type locality. 1. 1948.— Mertens. Peru). 23. 1803. pi. p. 1873a. BOLIVIA: Guajaramerin. p. Orellana. 99.— Aleman. 1802. 1824a. — — — — Rivero. p.— Barbour and Noble. JP 255.233 ft. 144673. PERU: Domo Santa Clara. locality. 145080. Bufo agua Daudin. 284. 1-3. p. USNM 127934-6. 29. 182. 1939. 7965. [37]. 12 km. 149713-21. 1824. 75. 481 (type locality. 1820. pi. p. 45. p.

nearly vertical and in a line with the upper lip. an adult male from Mitii. Colombia. in postaxillary region equal to greatest width of head. a median . and lower femur. no skinfold across the chest. a heavy skinfold on heel and knee. foot. broadly ovate. ending near heel. head width.S. side of head sepia. ending at parotoid. — Dimensions. When hind leg is adpressed. free. some of the tubercles brown mark between the clove brown. upper limb surfaces seal brown. abdomen.106 U. . knee and elbow are separated when hind legs are bent at right angles to body. side of the crest along upper lip in body vandyke brown fading with many dark sepia spots. much greater than interval between nostrils. triangular. both reaching beyond base of penultimate phalanx of third. their distance from end of snout about one-third their distance from eye. hand. 25. first finger longer than second and equal to fourth. external vocal sac in the male. prominent. 144619. Nostrils lateral. separated from eye by an interval equal to one-half its own diameter. strongly projecting. a very heavy tarsal ridge. a heavy glandular ridge from posterior corner of eye above tympanum. upper front of eye with seal lip brown wood brown. 97. interorbital diameter 1% times that of upper eyelid. round outer one. tibia. Skin of upper parts granular. soles granular. venter finely granular. a small semilunar metatarsal tubercle and a smaller flat. head length. a flat oval thumb pad present. the distal ones of the third and fourth fingers double. Parotoids very large. heels barely touch. venter clay color. with transverse folds on throat. bony areas between eyes and near crests nearly smooth. when limbs are laid along the sides. a large rounded palmar callus. partly encircled —Dorsum by Vandyke brown.. its diameter 1^ times its distance from tip of snout. with heavy sepia reticulations which become small and scattered on to clay color below. heel reaches nearly to shoulder. 28 mm. 36 mm. Color in alcohol. Heavy bony ridges on top of head. snout nearly truncate at tip when viewed from above. NATIONAL MUSEUM BULLETIN 2 88 — USNM Description. loreal region concave. Vaupes... 37 mm.. its greatest diameter one-half that of eye.5 mm. truncate in profile. Tympanum distinct.. Tongue about half as wide as mouth opening. 36. Toes moderately long and one-third webbed. femur. metacarpal tubercles well developed. dots. 38 mm. a row of large tubercles along the sides behind parotoid.5 mm. Fingers fairly long. a clove eyes. with indistinct darker marblings. with lateral ridges. twice the diameter of eye. with large spinose warts on each side of midline and toward the sides. its posterior border free and unnotched. Can thus rostralis strong.. its disk reaching halfway on antepenultimate phalanx of fourth. third toe longer than fifth. Eye large. Body stout. the upper jaw extending slightly beyond the lower.5 mm. Head and body. no ventral disk.

Colombian Specimens Examined Amazonas: USNM Leticia. p. Tolima: Melgar. 144617. 2 km. USNM 144614-6. soles of feet and palms of hands sepia. Vaupes: El Centro. 146818. 1953. USNM 144612. often has confused herpetologists. 148124-32. 279 (type locality. Bufo blombergi Myei-8 and Funkhouser Plate 17a-c 1951. USNM 13870. 1962. 56. Nachao. Lebrija. Quetame. color photograph. Narino. Cochran. USNM 148134. 147481. Copping. Gir6n. Females have lighter backs with a row of dark spots on either side of the middorsal region. Magdalena: Cienaga. 144585-6. Camp La Macarena. upper Rio Magdalena.— FROGS OF COLOMBIA — COCHRAN AND GOIN 107 lower proximal femur. A detailed study of Bnjo marinus from all the areas in which it occurs is needed before the possible subspecific status of the toads from certain localities can be verified. upper Guayabero. Guadalajara River near Santander: territory of Buga. USNM 145772. near mouth of Caiio Losada. USNM 144602-3. Norte de Santander: Catatumbo. Valle: Buenaventura. USNM 148133. with dark vermiculations. The identity of young individuals. Gallardo. upper Rio Giiejar. 96. 144605-8. 15-20 km. USNM 144587-601. and usually it is the most numerous species of the genus wherever it occurs. south of Cali. near Turbo. USNM 142206-8. 1957. USNM USNM USNM USNM USNM USNM 144613. The huge triangular parotoids make it readily recognizable. and its large size and enormous fecundity cause adults and young to be well known and frequently collected throughout its range. p. USNM 124174. USNM 148793. 146391-2. northeast of Villavicencio. Bufo blombergi Myers and Funkhouser. so far without much success. posterior seal brown fading below to clay color. Mitu. femur — Remarks. p. Meta: USNM 144154. 511. Choc6: Truando. Golfo de Uraba. SU 20305-8. Colombia). Antioquia: USNM MedeUin. 148121-3. Attempts to describe and validate local subspecies in South America have been made. p. USNM 12241 (juv. This extremely hardy and adaptable species is found over the whole of Colombia except in snow-covered regions. north of Camp Tibu. the tubercles and toe tips darker. USNM 144618-20. throat of male with black suffusions. 126860-3. 145078-9. 146390-1. — — — . Atlantico: Sabanilla. Rio Calima near C6rdoba. Villavicencio. with their much less developed parotoids and with the spotted coloration of the female. Narino: Rfo Satinga. BoLfvAR: Sincelejo. 144609-11. USNM 147250-1. Cundinamarca: Beltran.). Rio Apaporis. Cei. 1961a. USNM 144604. La Girona. 7 km.

ending near heel. a little more raised a heavy bony ridge from posterior corner of eye above tympanum.108 U. its diameter two-thirds its distance from tip of snout. blunt lateral ridges. sides of body dirty olive buff with darker tubercles. heels are slightly separated. first finger much longer than second. a faint skinfold across the chest. a poorly defined round palmer callus. a distinct skinfold on heel and knee. third toe much longer than fifth. another wide bony ridge from inner posterior corner of eye to nuchal region. Toes long. when limbs are laid along the sides. Dimensions. slightly projecting. oval. with faint darker reticulations and with the large blunt tubercles tipped with clove brown. heel fails to reach the shoulder. Can thus rostrails blunt but distinct. the upper jaw scarcely extending beyond the lower. its length equal to distance from tip of snout to posterior border of eye. soles of feet and palms of hands Description. 79 mm. 80 mm. when hind legs are bent at right angles to body. 80. hand. femur. loreal region concave. tibia. 212 mm. an adult from 50 miles north of Cali. merging gradually with the upper lip. blunt scapular ridges. oval metatarsal tubercle and a rounded outer one. posterior belly wood brown. a large blunt thumb pad present.. with very faint. Skin of upper parts coarsely glandular. 53... head length. NATIONAL MUSEUM BULLETIN 288 — USNM 142101. a pair of wide. sides of chest and anterior belly olive. a heavy glandular fold along side from parotoid to groin. their distance from end of snout about one-half their distance from eye. Nostrils lateral. Dorsum drab with very indefinite sepia mottlings. Body very stout. head width. Fingers moderate in length. 60 mm. its tip reaching to middle of antepenultimate phalanx of fourth. interorbital diameter a little over twice that of upper eyelid. oval. Eye large. a heavy. its greatest diameter a little over one-half that of eye. . onehalf webbed. knee and elbow are widely separated. vertical. blunt. — — . prominent. ending at anterior end of parotoid. throat.5 mm. the granules sepia. Valle. with many small tubercles on posterior part of back and sides. no ventral disk. 68 mm. Color in alcohol. snout broadly rounded when viewed from above.5 mm. truncate in profile. much greater interval between nostrils. Parotoid gland thick and prominent. with an occasional small olive buff spot. tongue half as wide as mouth opening.. free. separated from eye by an interval equal to its own diameter. those on throat and chest smaller.S. in postaxillary region wider than greatest width of head. venter granular. its posterior border free and unnotched. metacarpal tubercles well developed. Heavy bony ridges on top of head. fourth longer than either. an external vocal sac in male... sides of head dull olive with very indistinct darker markings. Colombia. Head and body. the granules on belly nearly flat. a distinct tarsal ridge. foot. When hind leg is adpressed. Tympanum not very distinct. upper limb surfaces olive.

p. separated from eye by an interval equal to one-third its own diameter. Supraorbital. and parietal bony ridges on top of head. .—Cope. postorbital. Choco.— Nieden. Rio Calima. foot. Bufo coniferus Cope. 1901. pi. 1898a. Specimens Examined COLOMBIA USNM Valle: 50 mi. 1962. sloping a little to the upper lip. near Anchicaya. p. p. tibia. 1862b. 96. p. p. New Granada [-Choc6]. 394. Fingers quite long. p. USNM USNM Bufo coniferus Cope Plate 16a-c 1862. 1932. Chilophryne conifera 1932 p. near Guaduas. 1946. p. p. fig. hand. 1923. slightly with lateral ridges. 321.—COCHRAN FROGS OF COLOMBIA wood brown. 1862a. Turbo region. strongly projecting. Colombia). 1899.— Gallardo. 50. El Bargell. nearly truncate in profile. 358.— Nieden. their distance from end of snout one-half their distance from eye. 43. slightly concave. 65. p. Colombia) . north of Cali. 29. loreal region vertical. 145775.— Kellogg. fourth longer than . 1882a. tips of toes and AND GOIN fingers darker. 102. concave in outline. Canthus rostralis very strongly ridged. 123. has the following millimeter measurements: head and body. p. 1863.— Kellogg. The critical proportions are very close to those of the described specimen. B. 50. head width.— Kellogg. 1923. Tympanum distinct. Lucilius coniferus. — USNM Description. 251. 133. interorbital diameter twice that of upper eyelid and of interval between nostrils. snout widened at tip. Bufo ehlersi Werner. 29. 158 (type locality. p. 29. p. 1865b. 481 (type locality.— Boulenger.5. femur. 172. Colombia. 66. The only important differences are that the tympanum is much more distinct in the smaller specimen but is only two-fifths the diameter of the eye and the toes are about one-fourth webbed. Tongue about two-fifths as wide as mouth opening. p. 69. 60. with a small vertical ridge when viewed from above. Eye large.— Giinther. 102.— Breder. — 109 webs between toes USNM 142102. an adult from Turbo. 133. p. prominent. head length. Nostrils superolateral. p. spatulate. p. 4335 (holotype). its posterior border free and unnotched. 145774. the two first continuous in an obtuse angle. 1865b. its diameter equal to its distance from tip of snout. 1932. 63. its greatest diameter one-half that of eye. 1914. 142101-2. the also wood brown. the upper jaw extending well beyond the lower. with the same data as the described specimen. 1863. Another specimen at hand. 1862. first finger webbed and shorter than second. a female Remarks. 813. —Cope.

its tip reaching triangular thumb pad halfway along antepenultimate phalanx of fourth. 27. no ventral disk. venter uniform russet.3. articulation well — Dimensions. when limbs are laid along the sides.S. Turbo. knee and elbow overlap.4. its tip reaching to base of penultimate phalanx of third. Head and body. the head between supraorbital ridges flat and nearly smooth. when hind legs are bent at right angles to body.. oval inner metatarsal tubercle and a smaller round outer one. Body stout. Toes moderately long. tibia. 75 mm. a short bony ridge from posterior corner of eye above tympanum. side of head chocolate with a pale cinnamon diagonal bar from lower eyelid to corner of mouth. a small present. metacarpal tubercles well developed. a large squarish palmar callus. the digital tips and Color in alcohol. NATIONAL MUSEUM BULLETIN 2 88 either. When hind leg is adpressed. limbs with wide. ending at parotoid.5 mm.29. mm. a pair of crescentic pale cinnamon spots on shoulders just behind parotoids and another pair on sacrum.. —Dorsum cinnamon with suggestions of large.27. parotoids small but with several large warts. 1901.5 mm.5 . sides of body chocolate lightening to cinnamon below.89.58. a median external vocal sac in male. on either side of the paler midline. a small. PANAMA: BM BM 1914. with scattered warts from back of head to above anus.28. third toe slightly longer than fifth. hand 26.142 from Cachabe. BM USNM 4335 (type).6. 98. with darker markings in front of and behind it. Skin of upper parts granular. femur.110 U. foot. becoming paler on belly.21. Ecuador. BM USNM 65124. heels touch. Cayapas River. 32.5 mm. 1901.5.5 mm. Specimens Examined COLOMBIA Choc6: Peiia Lisa. inconspicuous dark bands above. head length. San Javier. chocolate spots tubercles a little lighter. ECUADOR: Cachabe. . a heavy skinfold on heel and knee. Condoto.. a row of spiny warts from posterior end of parotoids along sides to groin.17-9.7. venter granular with small warts on side of belly and below thighs. palms of hands and soles of feet wood brown. Pambelas. single.28. head width. The prominent nostrils and small projecting vertical ridge on the snout may suggest a slight relationship with Bujo granulosus. measures 90 millimeters in head and body length. — BM Remarks.. USNM 53719. while the toes are one-half webbed. BM 98. 30. heel reaches posterior corner of eye and tarsometatarsal beyond snout. The first finger is appreciably shorter than the second in all.4.29. onehalf webbed. in postaxillary region slightly less than greatest width of head. Porto Bello. no tarsal ridge..33-4. upper Trinidad River.5 mm.141-5. 31. a slight skinfold across the chest. 22. The largest of 16 examples at hand. 1902. irregular.

Puerto Cabello. pp. Fingers moderately long. 8. p. a large. 311. 4 Kellogg. p. its greatest diameter one-half that of eye. reaching to base of penultimate phalanx of fourth toe. with a few larger ones intermingled with them. High sharp. 68.— Rivero. rather indistinct end of 'Two cotypos from Cordoba. 1919. a bony postorbital ridge from posterior corner of eye to tympanum. p. Miiller. 1914. conical tubercles from end of postorbital crest over middle of parotoid and along the side diagonally downwards to groin. — Shreve. 352. Nostrils dorsolateral. 535. 96. 1868. p. 1947a. 1882a. but truncate at tip when viewed from above.1.— Boulenger. a similar line of pointed tubercles along outside of foot from heel to end of fifth toe. the upper jaw extending considerably beyond the lower. curving ridge. distinctly projecting. 68. when limbs are laid along the sides. Body stout. pi. 278. Cundinamarca. fourth longer than either. 1923. p. — — — p. its posterior border free and unnotched. 45— Gines. p. prominent. 1964b. vertical. 1901. p. 1961. 279 Peracca. Skin of upper parts covered with small tubercles. a line of pointed. p. a distinct skinfold on heel and knee. a short. p. its tip reaching to base of penultimate phalanx of third. 294. with lateral ridges. 323. their distance from snout one-half their distance from eye. its diameter equal to its distance from tip of snout. 5. snout with a narrow. 1959.15. Toes fairly long. c (type locality. . free.FROGS OF COLOMBIA — COCHRAN AND GOIN 111 Bufo sternosignalus Gunther Plate 13d-e 1858. fig. 247. merging with the upper lip. heel reaches posterior corner of eye. p. loreal region concave. third and fifth toes subequal. Bufo sternosignalus Gunther (part*). a small but prominent oval metatarsal tubercle and a smaller round outer one.— A. p. 1867.9. p. p. vertical ridge down the front. 1927. 3. Tongue half as wide as mouth opening. soles granular. with a few larger granules in front of anus. Colombia. interorbital diameter twice that of upper eyelid and twice interval between nostrils. Venezuela).— Smith and Taylor. 1884. When hind leg is adpressed. 108. 38. Bufo typhonius sternosignatus. first finger a little shorter than second. Lutz. Eye large. pp. 1932.— F. 70— Mello-Leitao. separated from eye by a very minute interval. 28. round palmar callus. pi. 1937. somewhat elevated above head surfaces. 42. 118. one-half webbed. a small but prominent oval thumb pad present. Tympanum distinct. 1916. bony ridges on top of head. Mexico.— Beebe. are now considered to be Bufo valliceps Wiegrnaun.— Werner.— Keferstein. in postaxillary region equal to greatest width of head. oval. a young male from Bogota. p. 1962. Canthus rostralis bordered by a sharp. p. 132. p. —BM Description. no tarsal ridge. 1947. Parotoids small. metacarpal tubercles well developed.— Gallardo. 1948. 207 Nieden. when hind legs are bent at right angles to body. figs. slightly rounded in profile. heels touch. venter finely granular. knee and elbow touch. 68. 13. 302.

head width. with small sepia spots on throat and anterior Color in alcohol. 16... Head and body. a pale middorsal line from snout to anus.: 112 U. and especially those behind the ear and along the side. and Boulenger. tibia.1-2 Andes of Venezuela.5 mm.. 15 mm. In typhonius.S. BM 94. Remarks. But in coniferus the parietal crest reaches a much greater development than it does in the other two. 1882 (Bogota). NATIONAL MUSEUM BULLETIN 288 parietal ridge. On the strength of Boulenger's identification of the Bogota frog.31. head length. typhonius in having a short vertical ridge along the tip of the snout. p. upper limb surfaces ochraceous with a few wide sepia crossbars.9. 535) assigned sternosignatus a subspecific rank under typhonius.5 mm. Dimensions. are much higher and more "spiky. These records probably refer to B. hand. in coniferus at least. foot 13. 1820 M. p. — Dorsum raw sienna. venter buff..15. the largest one below anterior corner of eye. Angelopolis. BM 68. Shreve (1947a. a large sepia patch from corner of mouth to postocular ridge. a distinct skinfold across the chest. and other indistinct dark diagonal markings on the back. 1914 (Cafetal Camelia. VENEZUELA: . with three or four sepia spots on upper lip. abdomen and a larger triangular sepia mark on center of chest head ochraceous-buff.8. Dunn (1944) does not mention this species in his "Herpetology of the Bogota area. apparently does not increase in older and larger animals to the fantastic proportions found in typhonius of similar size. which.5 disk. while in sternosignatus the jaws are normal." but he includes Bufo typhonius in the actual fauna of Bogota . femur. this spciees is retained in the Colombian fauna. 39) discredits the presence of B. palms of hands buff. typhonius alatus. — mm. inner half of sole of foot buff. All three likewise have a raised postorbital crest. Werner.. sternosignatus in Colombia as follows half of side of — Reported from Colombia by Peracca.12.. covering tympanum. nr. a weak ventral an external median vocal sac in the male.). 14.. the outer half pale sepia. 40 mm. 11 mm. Specimens Examined COLOMBIA Cundinamarca: Bogota.. 14 mm. 1916 (Canon de Tolima). without forming a right angle. This species resembles Bufo coniferus and B. .. a dark chevron-shaped mark on upper eyelids and interocular region." Both typhonius and sternosignatus have a curving crest around the eye merging gradually with the parietal crest. the angle of the jaws is produced beyond the usual head outline. in fact. making nearly a right angle with the subocular crests while the tubercles on the back. and extending backwards below the line of pointed tubercles on parotoid and along the side through groin and along anterior femur. Rivero (1961.

reaching halfway on penultimate phalanx of third. — 10.— —COCHRAN FROGS OF COLOMBIA AND GOIN 113 Bufo glaberimus Gtinther Plate 16d-p 1869. Eye large. one-half webbed. Grande. their distance from end of snout about one-half their distance from eye. 1882a. an external vocal sac in the male? Dimensions. 12 mm. truncate in profile. 16 mm.5 mm.. its diameter 1% times its distance from tip of snout. Peracca. 13 mm. tongue half as wide as mouth opening. No bony ridges on top of head. Can thus rostralis distinct. 2 (type locality. Body stout. Curtdinamarca. p. nearly vertical. all single. with a few minute pustules between the eyes and some faint tubercles on sacrum. 483. slightly projecting. 37. loreal region flat. with a heavy lateral skinfold from the posterior outer border along sides almost to groin.. heels slightly overlap. . third toe much longer than fifth. head length. ending near heel. foot. a small but prominant oval inner metatarsal tubercle and a smaller round outer one. p. p. Colombia.— Gallardo. obovate. 1923. Description. pi. When hind leg is adpressed. Parotoids large. separated from eye by an interval equal to one-fourth its own diameter. 96. Toes moderately long. fig. free. 1% times the eye diameter. 14. knee and elbow are narrowly separated. interorbital diameter three-fourths that of upper eyelid. its greatest diameter two-fifths that of eye. slightly concave before merging with the upper lip. a distinct ventral disk. — — USNM 144566.— Nieden. subarticular tubercles distinct. ending at parotoid. Boulenger.. its posterior border free and unnotched. rounded palmar callus. snout rounded when viewed from above. its disk reaching halfway on antepenultimate phalanx of fourth. 1962. prominent. equal to interval between nostrils. head width. a faint skinfold across the chest. oval. p.5 mm.5 mm. a half-grown specimen from Cano Meta. 112. lower leg surfaces more heavily granular. Head and body. p. throat and chest nearly smooth. Colombia. 1898a. heel reaches to shoulder and tarsometatarsal joint reaches to end of snout. femur. heavy glandular ridge from posterior corner of eye above tympanum. with lateral ridges. when hind legs are bent at right angles to body. a large. when limbs are laid along the sides. a heavy skinfold on heel and knee. 37 mm. 14. hand. Bufo glaberrimus Gunther. in postaxillary region equal to greatest width of head.. a blunt tarsal ridge. a short.. 123. Tympanum distinct. a small round thumb pad present. the upper jaw extending slightly beyond the lower. fourth longer than either. nearly longitudinal. 1904. Villavicencio. metacarpal tubercles well developed.. 291. Fingers moderately long. Nostrils lateral. p. 35. belly faintly granular. tibia. first finger a little longer than second. Bogota. Skin of upper parts nearly smooth (glandular under lens).

5. p. 1868. 123. groin buff with a vertical sepia spot in front of it.12. Description. an adult female from the Virology Field Station. The ventral region is pale or faintly spotted in young specimens. pi. p. 1867. BM 88. p. p.5.— Peracca 1904. upper leg surfaces with indistinct traces of two or three darkedged crossbars. p. Villavicencio. the toe tips and subarticular tubercles buff. Rio Raposo. 35. the largest of which are concentrated along each side of the midline.— Boulenger.123-8. region of Truando> Colombia). Meta: Cafio Grande. 9. In specimens over 50 millimeter long. becoming drab gray on belly. ECUADOR: Cachabe\ BM 98. 157 (type locality. Vattpes: Amanaven. p. p.— Nieden. BM 80. posterior femur and tibia with smaller sepia reticulations on a buff ground. 147516. Sarayacu. 1876. soles of feet slategray to dull olive-buff. BM 80. boca del Rio Giiaviare. —Dorsum immaculate smoke-gray parotoids wood brown. 353. 290. BolIvak: Alto de Quimari. 94. p. a wide seal brown lateral stripe beginning behind tympanum on outer third of parotoid.129-31. 1862a. 243. 393 Bufo haemalilicus Cope. side of head olive-gray with a few traces of sepia spots below eye and along lip border. 1863. 1913.178. Keferstein. — Specimens Examined COLOMBIA CNHM 61767-9. p. upper Rio Giiejar and El Mico. 3.28.16.114 U. palms of hands similar but lighter. MLS 63(a-d). 1862b. p. Colombia. 110. BM 95. 1923.228. 15.28.4.12. but it often becomes clove brown with scattered pale dots in the large. the back is usually uniform light brown or gray. 1962. Rhaebo haematiticus—Cope. Valle: Buenaventura.11. No bony ridges on top of . 1022— Giinther. — USNM p. Some are almost white (in alcohol) with black markings.— Andersson. This toad seems less brightly colored than other specimens of nearly the same size or smaller. USNM 144569-70. Remarks. Cundinamarca: Bufo haematiticus Cope Plate 16g-i 1862. USNM 144556-64. New Granada — [Choco].62-3. NATIONAL MUSEUM BULLETIN 288 Color in alcohol. p. Macarena. Villavicencio USNM 144566-8. smaller black markings thickly cover the outer parts of the back. lower leg surfaces coarsely mottled with cream buff and drab gray. Paramba. 1939. p. p.4. 1882a.— Breder. 49. 1946. — Gallardo. the first finger is usually longer than the second. although in a few examples these two fingers are subequal. USNM 71121. Valle. 1901.S. arm olive-buff with a few pale sepia reticulations beneath and a darker reticulation on elbow. 1862. The toes are one-third to one-half webbed. 357. old individuals. fig. continuing below lateral skinfold and fading out midway to groin. p. throat and chest cream buff. anterior femur heavily marked with long sepia bands on a buff ground. Bogota. 1898a. 96. 295.

palms of hands and soles of feet seal brown. a prominent oval inner metatarsal tubercle and a smaller round outer one. Tympanum distinct. but with fine folds across throat and most abdomen. interorbital diameter 1% times that of upper eyelid. 20 mm. single. vertical. Tongue about — COCHRAN two-fifths as wide as AND GOIN mouth 115 opening. heel reaches center of eye and tarsometatarsal articulation extends far beyond tip of snout. free. with very weak lateral ridges. when hind legs are bent at right angles to body. upper limb surfaces with indistinct darker crossbands. fourth longer than either. its tip reaching halfway on antepenultimate phalanx of fourth. third toe a little longer than fifth.FROGS OF COLOMBIA head. venter smooth. Skin of upper parts weakly tubercular everywhere.. 15. heels overlap. no skinfold across the chest. projecting. with a paler spot from anterior corner of border. declivitous in profile. 16.5 mm. Canthus rostralis sharp and straight.distance from end of snout one-half their distance from eye.. When hind leg is adpressed. an external vocal sac in male. in postaxillary region narrower than greatest width of head. Head and body. knee and elbow slightly overlap.. prominent. Body elongate. Color in alcohol. 12 mm. femur.. head length. its posterior border free a rather indistinct tarsal ridge. spatulate. its diameter equal to its distance from tip of snout. 22 mm. a series of skinfolds on heel and knee. side of body sepia. when limbs are laid along the sides. its tip reaching to base of penultimate phalanx of third. —Dorsum sepia. Fingers rather long. its greatest diameter three-fifths that of eye. venter seal brown anteriorly. mottled with fawn color below. and uiniotched. metacarpal tubercles well developed. 46 mm. a small oval thumb pad present. first finger much longer than second. separated from eye by an internal equal to one-fourth its own diameter. of corner of eye above short. the posterior part of belly which is and lower surface of limbs fawn brown spots except on proximal lower femur unspotted. even between eyes and on eyelids. with a pair of clove brown spots between shoulders and another pah* just anterior to sacrum. Eye large. foot. color with heavy seal head clove brown.. which is as long as its distance from nostril. a narrow glandular ridge from posterior parotoid. ending near heel. 15 mm. posterior femur mottled sepia and fawn color. then. almost twice the interval between nostrils.. Nostrils lateral. the upper jaw extending well beyond the lower. no ventral disk. loreal region flat. . Toes fairly long. a larger squarish palmar callus. merging with the — Dimensions. tympanum. tibia. one-fourth webbed.5 mm. descending to the upper lip. band. the tubercles a little side of eye to lip lighter. head width. snout broadly rounded when viewed from above.

oval. Colombia).5. snout broadly rounded when viewed from above. Antioquia. Body moderately stout. fig.11. pi. Truando. USNM 145077. Andagoya. without lateral ridges. Rfo Raposo. inner metatarsal tubercle and a smaller round outer one. Condoto. their tips reaching to base of antepenultimate phalanx of fourth toe. an indistinct. its posterior border free and unnotched. their distance from end of snout about one-third their distance from eye. Rio San Juan. its diameter 1% times distance from tip of snout. heel reaches poster- indistinct tarsal ridge. opening. apparently ending at heel. their length equal to distance tongue two-third as wide as from mouth tip of snout to center of eye. third and fifth toes subequal. Juntas. Parotoids large. Since the co types are discolored and in poor condition more than a century. ridges on top of head. Bufo hypomelas Boulenger p. in one-third BM webbed The toes are from one-fourth to examples at hand.21. prominent.12. finger shorter heavy skinfold on and knee.87-8.57-61. ovate. an adult from Andes. Eye large. equal to interval between nostrils. The maximum known head and body length is 90 after millimeters. 124233. When hind leg is adpressed. 2 88 — Remarks.16. Catatumbo.69-70. Santander: Rfonegro. distinctly projecting. fourth longer than either. Colombia. BM USNM USNM 48448-9 (cotypes). 4344. subarticular tubercles not strongly developed.11. Virology Field Station. Can thus rostralis distinct.11. and the tarsometatarsal 1914. Bufo hypomelas Boulenger Plate 17d-f 1913. Condoto. vertical.S. Colombian Specimens Examined BM USNM 1910. in all articulation extends far beyond the tip of the snout.88. Norte de Santander: Camp 15-20 km. west of Playa de Oro. I (type locality. metacarpal tubercles well developed. 147515-6. truncate and sloping backwards in profile. Description. the upper jaw extending well beyond the lower. the upper lip flaring out below it. 1913.103-6. north of Tibu.21. free. loreal region flat. Choc6: Pefia Lisa. 1022. Toes rather short. one-half webbed. Choc6. 147148-58. a small oval thumb pad present. 1914. in postaxillary region equal to greatest width of head.5. first than second. Valle: Buenaventura. elongate oval. a heel . 144679-80. interorbital diameter a trifle greater than that of upper eyelid. a fresh example has been chosen for describing and figuring. 102. USNM MLS 249. —AMNH No bony 14031. its tip reaching beyond base of penultimate phalanx of third. BM 95. 10-15 km. a large round palmar callus.7. a very small. Nostrils lateral. Tym- its panum Fingers long.116 NATIONAL MUSEUM BULLETIN U.

7.5 foot. with a few small drab spots. B. 9. hypomelas identify. the type of blombergi is 207 millimeters long. 9 mm. with paler metacarpal and metatarsal tubercles. with some faint tubercles on head and upper eyelids. tibia. mm. — In structure.. (Length of the type specimen was said to — Dorsum Vandyke brown. and they represent the midget and the giant of that group. the first of these extending onto eyelids. femur.) . The tympanum of the former is indistinct. hand. heels slightly overlap. sides of head wood brown. about 1% times the length and oneior touch. 337-262—70 9 makes it one of the easiest toads to . traces of a pale longitudinal line along dorsolateral region. elongate parotoid gland. with several larger darker spots along upper lip and in loreal region. The heel of AMNH 14031 reaches the posterior border of the eye. in the latter it is moderand equal to one-half the eye diameter. The very distinctive color pattern of B. rounded. 10 mm. With but in critical a single specimen at our disposal. an external vocal sac in the male? half the — Head and body. head length to angle of mm. the range of variation measurements remains unknown.. Both of B. sides of body mostly seal brown. Skin of upper parts minutely pustular. light bordered spots on either side of midline. no ventral disk. lacks the parietal crests of blombergi (such young specimens) and the than the second in hypomelas but is first longer species obviously are in the "guttatus" group. 7. knee and elbow when hind legs are bent at right angles to body. webs between toes pale wood brown. hypomelas is only 23. a short but heavy postocular crest. with a few granules around the vent. venter drab with heavy sepia reticulations that are closer together anteriorly.5 mm. Color in alcohol. These differences might be bridged if more material were available. upper limb surfaces pale russet with wide seal brown crossbars.. when limbs are laid along the sides. venter smooth. the last more chevron-shaped and just behind the sacrum. head width. blombergi except that it Bufo hypomelas resembles the young crests are not always developed in finger is slightly shorter in blombergi. 23. width of the eye..FROGS OF COLOMBIA —COCHRAN AND GOIN 117 border of eye.5 mm. no skinfold across the chest. Dimensions. followed by the narrow. soles of feet and palms of hands drab. jaw. 7.5 millimeters long. while that of the type was said to extend to the tip of the snout. with several large.. Remarks. The color pattern 14031 where it remains is like that shown in the figured of ately distinct — AMNH — type specimen.5 be 20 millimeters.5 mm.

metacarpal d 1 . projecting beyond lower. c1 . dark -spotted posteriorly. venter pale. entire. pedimarmoratus Heel reaching to posterior corner of eye. h size to 37. immaculate. mm . Fingers free or at base. A. A. Diapobehind. l g /'. size to 23. ebenoides marinkellei mm Belly immaculate cream. disk. Metacarpal tubercles absent or weak. varius elegans Metacarpal tubercles well developed. e1 . color mainly black. Tongue No tympanic elliptical. spurrelli mm Venter immaculate. l . varius subornatus Belly dark with a few large light spots. Fourth toe long. size to 28. Outer metatarsals united. tubercles well developed... fairly smooth above. Belly slate color with large light paired spots. Atelopus Dumeril and Bibron. connected by the epicoracoid cartilage.. free webbed more or less webbed. Family Atelopodidae Girdle firmisternal. mm . ignescens Skin thinner. or scarcely.5 . g 2 mm P. 660 (type species. A. 1 ft . —Pupil horizontal.5 . size 30 developed. A. Skin thickened. longibrachius webbed. A. No omosternum. . size to A. nicefori Heel reaching between center of eye and nostril. throat with a dark spot. Eggs Genus Atelopus Dumeril and Bibron 1841. ebenoides ebenoides Upper jaw extending appreciably beyond lower. Terminal phalanges simple. Key a a2 1 . flavescens Dumeril and Bibron). pachydermus Arms and legs not unusually large. 2 . Eye diameter equal to or slightly greater than snout length. /'. Belly light anteriorly. toes nearly completely A. laid in water. Sacral diapophyses dilated. Upper jaw not. Tadpoles aquatic. A. All toes normally tympanum 48 b2 to . mm . often rough and tubercular. c1 . Generic diagnosis. sternum cartilaginous. Heel not reaching beyond posterior corner of eye.5 A. Colombian Species of Atelopus Arms and legs massively developed. toes one-half webbed. throat and chest with dark longitudinal spots. toes physes of sacral vertebra moderately dilated. . one or more digits sometimes lacking.118 NATIONAL MUSEUM BULLETIN 288 U.5 e1 . other toes poorly developed and relatively short. c2.S. size to 25 A.5 mm. mm hidden. p. size to 45.. size to 32. Coracoids and precoracoids more or less divergent. Palate smooth. d2 . bufoniformis tympanum distinct mm anteriorly. Heel reaching center of eye or beyond. size to 50 A. size to 38 g e2. l . g Belly light. h2 e2 .. size to 53 mm . A. mm. the tips not dilated into regular disks. no omosternum. Heel not reaching beyond front of shoulder or angle of jaw.

outer ones one-half webbed. 3. mm c2. p. size to 33 Eye diameter less than snout length. belly pale. Redrawn by Paul Laessle after O. heel reaching d between axilla and posterior border of eye. immaculate. larger ones on belly. Schmidt. metacarpal tubercles absent. 84. /'. size to 44 A. heel reaching posterior corner of eye. p. 1 . Nieden has synonymized this species with Atelopus cruciger. Hohe von 5000' "). — Rivero. pi. e1 mm A. p. but Venezuelan examples of cruciger have much smaller feet and hands and a dorsal pattern different from that shown in Schmidt's figures. A. walkeri . smooth. carrikeri Skin thinner. fig. pustular. 1963. inner toes entirely. 26 (type locality. 1926. Atelopus cruciger (not of Lichtenstein and Martins). 1963c. longirostris e2 Metacarpal tubercles well developed.— FROGS OF COLOMBIA — COCHRAN AND GOIN 119 Heel reaching angle of jaw to posterior corner of eye. "westen Bonaventura [= Buenaventura. belly slate gray. Skin thickened. size to 46. Metacarpal tubercles absent. Schmidt (1858. approximately size). 107. chin and chest dark. von Neu-Granada. toes one-half webbed. small dark spots on chest. mm Atelopus pachydermus (O. varius glyphus f2 Heel reaching center of eye. 1926. toes one-half webbed. 257. p. toes one-half webbed.5 mm. Heel reaching to posterior corner of eye.5 d2 . Phirix pachydermus O.[telopus] pachydermus. Valle] in einer — Nieden. size to 51. fig. Schmidt) Figure 6 1858. varius bibroni half webbed. belly pale. Atelopus pachydermus (type of Phirix pachydermus. size to 39 A. bei pi. toes oneA. . immaculate. 3. chin with a light spot below angle of mouth. A. Figure same 6. mm . 26). .

82. an adult female from Paramo de las Papas. The skin texture. Cauca. Description. snout short. a rudiment thumb [metacarpus] distinct and likewise a distinct although very flat The of the tubercle on the outer ball [palm] besides. and indeed the remaining toes do not possess the usual mobility. on the femur. color sulphur-yellow on whitish. the depth of the occiput is bordered by a pair of swollen borders. truncate in profile. The sole appears very broad and solid.S. Head as long as broad. p. prominent snout tip as from the inner eyelid. p. Atelopus bufoniformis Peracca. NATIONAL MUSEUM BULLETIN 288 Until adequate material has been collected near Buenaventura. 107. A distinct mention is merited of the blackish color of Color: which in the top of the first finger of the forelimb. 75 mm. with blue lines on the back. shows this thickened characteristic. The original description (O. forelimb. p. 1858. in fact. leathery. The back shows. a pair of continuations of these blue lines are extended on the thigh. the name pachydermus is provisionally retained. snout prominent. 1963. continuous irregular blue markings the center enclose three light spots. Size: Habitat: of Atelopus bufoniformis Peracca Plate 18a-c 1904. Their web is so thick and. Colombia. On the tarsus of the hind foot are two weak tubercles. rather regular and symmetrical. the first and second so that they appear only as a pair of blunt tubercles. Ecuador) Nieden. 24. The nostrils are almost as far from the rounded [dome-shaped].— Rivero.— 120 U. [Valle. although of medium size. As we have already mentioned under generic characters. and fifth toes are buried in the connecting web. Its upper central part is horizontal and deepened. 53 mm. broadly angulate when viewed from above. forelimbs are unusually stout. Western New Granada. first finger thick.. —MLS 143.] at an elevation 5. short. and towards the second finger. the head. Very rare. 257) is translated as follows: [Diagnosis:] A Phirix with both anterior and posterior feet very robust. and also on the posterior femur. which appear as the continuation of the equally swollen supraorbital borders. p. Tongue as wide as mouth opening. Nostrils two-fifths lateral. its posterior border free and unnotched. p. The webbing forms a border on the fourth toe from the level of the second articulation. head moderate. 40 mm. thick. It is so stiff between the first three toes that these cannot be laid side by side. Puno. hind limb. third. The entire venter is yellow without interruption. the hands indeed as if stuffed. and rigid. on a yellow ground. runs transversely over the posterior part of the back and on the underside of the thigh. is thick and marked with bending a little weals. Schmidt. Another blackish blue mark. near Buenaventura. The first. 1963c. Body. skin calloused. 1926. the upper jaw not extending beyond the lower. not projecting.— Gorham. oval. their distance from end of snout about one-half . this animal has a massive build in its flanks and limbs.. second.000 feet. resembling that of a pachyderm. 20 (type locality. that it gives scarcely the impression of being a web. so to speak.

femur. which is approximately equal to fourth. those on toes appearing to be arranged between in pairs. The snout from Puno. 9 mm. when hind legs are bent at right angles to body. Can thus rostralis distinct.. a is adpressed. flaring to the upper lip. — Dorsum seal brown anteriorly. —In most characteristics MLS 143 resembles the de- scription of the type of Atelopus bufonijormis Peracca Ecuador. Tympanum hidden. venter and sides with very coarse. 6. the largest warts in a fairly distinct row along the dorsolateral line.. when limbs are laid along the sides. no true stout. head length. with a row of smaller tubercles on each side of the midline. 9. Remarks. equal to the diameter of eye in the former. Fingers short.5 mm. heel reaches barely three-fourths the distance from groin to axilla. postaxillary region equal to greatest width of head. sole of foot entirely covered with coarse irregular granules. its diameter equal to its distance from tip of snout. no skinfold across the chest. third toe shorter than fifth. as in — mm. a little longer than that in the latter. 9.5 mm.FROGS OF COLOMBIA — COCHRAN AND GOIN 121 their distance from eye. a small oval thumb pad and a large round palmar callus present. metacarpal tubercles not distinguishable from the coarse granules covering entire palm of hand. 9. sides of head clove brown. the latter appearing very long in comparison with the others. 30 mm. Eye fairly large and prominent. no ventral disk. hand.. knee and elbow are widely separated. interorbital diameter equal to that of upper eyelid. first finger shorter than second. the tubercles a posteriorly. foot. a short glandular ridge from posterior corner of eye straight backwards above tympanic area and ending above arm insertion. loreal region concave. head width. fully webbed. First toe distinct. Dimensions. Head and body. the web reaching almost to its tip as a wide lateral ridge. palms and soles seal brown. with somewhat finer granules on limbs and throat. 9 mm. Skin of upper parts heavily tubercular and warty. the large round warts walnut brown. Body When heavy skinfold on heel and knee. a little less than interval nostrils. Color in alcohol. while No is it is mention is made in the original description of bufonijormis of . a small oval metatarsal tubercle and a small round outer one.. both reaching to base of penultimate phalanx of third.5 mm.. immaculate. vertical. trifle paler. while it is said to be almost double in the latter. large round warts. clove brown some of the warts a little lighter. venter and sides seal to clove brown. The most noteworthy difference is that the interorbital diameter equals that of the upper eyelid in the former. which reaches to center of antepenultimate phalanx of fourth.. tibia.5 lacks external vocal sacs. heels fail to meet. in hind leg tarsal ridge. The male probably has the forearm enlarged but most other species of the genus.

sloping steeply outwards to the upper lip. figs. and posterior femur. extending onto the sides and becoming larger. scarcely projecting. Colombia. 9 (type locality. its greatest diameter about two-fifths that of eye. in the adpressed arm. Huila. loreal region deeply concave. ending near heel. 3. heels are separated.. any reduction toe 4 NATIONAL MUSEUM BULLETIN in toes 1. Paramo de las interval equal to three-fourths its long. Toes rather short. equal to interval between nostrils. 1963c. San Agustin. 3. metacarpal tubercles weakly developed. 3. fourth longer than either and reaching to base of last phalanx of third. an adult female from Papas 3. heel reaches front of shoulder. the tips of all shorter than second. chest. digits bulbous. Tympanum indistinct posteriorly. third and fifth toes subequal. a very faint tarsal ridge. first finger own diameter. Head as long as broad. in postaxillary region equal to When hind leg is adpressed. venter pustular and slightly granular. when hind legs are bent at right angles to body. a small oval inner metatarsal tubercle and a smaller round outer one.S.600 m. Skin of upper parts thick. the upper j aw extending only slightly beyond the lower. pi.. their disks reaching to center of antepenultimate phalanx of fourth toe. described specimen was the only one examined. 2 88 with the resulting prominence of are exceedingly obvious in the Cauca frog.122 U. truncate in profile. Atelopus ebenoides ebenoides Rivero Plate 18d-f 1963.600 m. tip of third finger just reaches to end of body. — characters which The and 5. —CNHM 69746 (holotype). 3. with thick lateral ridges. its length a little more than one-fourth that of head and body. a large round palmar callus. Atelopus ebenoides Rivero. with many low round warts from level of shoulders backwards. its diameter nearly as great as its distance from tip of snout. its posterior border free and unnotched. a pair of bony arc- greatest width of head. Nostrils lateral. . a flat oval thumb pad present. separated from eye by an Description. 2. with small tubercles on throat. Body rather stout. a heavy skinfold on heel and knee. when limbs are laid along the sides. Colombia). snout nearly truncate when viewed from above. tongue very long and narrow. interorbital diameter V/2 times that of upper eye lid. knee and elbow are slightly separated. their distance from end of snout about one-half their distance from eye. prominent. San Agustm. one-third as wide as mouth opening. Eye large. a heavy bony ridge from posterior corner of eye extending straight back and ending in a small knob behind the tympanum. Paramo de las Papas. 6. p. Canthus rostralis rounded but prominent. Fingers moderately webbed much at the base. excepting the distal part of fourth toe. Huila. nearly completely webbed.

FROGS OF COLOMBIA — COCHRAN AND GOIN 123 shaped ridges behind the head extending onto anterior part of back.. side of head slate-black. 17 mm. series of of the large tubercles. bujonijormis the fourth toe USNM well with the holotype. tibia. same data as holotype.. It is 33 millimeters in length. mm. with paired buff spots below arm and on side of belly.5 mm. 151325. collected by Hno.. 48 mm.5 — Dorsum slate-black.. and the smaller webs between its toes. 150645-6. Boyacd. 16. Remarks. An interesting fact is that one of the two known examples of Colombia bujonijormis also comes from Paramo de las Papas. and with some median buff spots on posterior part of belly. Description oj holotype. An additional example from Tolima. 17. 13 mm. projecting upper jaw.third as wide as mouth opening. head length. rounded when viewed from above. an adult from P&ramo Vijagual.. head — width. CNHM 69746 (holotype). Color in alcohol. USNM 151325. with a dorsolateral round buff spots covering some mm. The feet are entirely — however. Tolima: Near Manizales on old road to Nevada de Ruiz. Atelopus ebenoides ebenoides resembles A. while in bujonijormis it is hidden under the thick.. 150644. bujonijormis chiefly in its truncate snout and its dark color. External vocal sacs in the male? Dimensions. Atelopus ebenoides marinkellei. no ventral disk. no skinfold across the chest. rounded in profile. more posterior nostril. with a pair of large irregular buff spots on either side of the chin. foot.600 m. new subspecies Plate 18g-i — USNM — USNM — Holotype. elongate. Head and body. its posterior border free and unnotched. the digits buff below. Diagnosis. soles of feet and palms of hands dull drab with a few buff spots. the upper jaw extending well beyond the — . in ebenoides the tympanum is distinct anteriorly. a small buff crescent below eye and a light dot below nostril. while in is relatively very long while the others are extremely short and poorly developed. venter slate- black. the type locality of ebenoides. 12 mm. and on upper and lower surfaces of tarsus. oval. Also. Resembles Atelopus ebenoides ebenoides except for its longer snout. tongue one. San Agustin. femur. Colombia.. as ebenoides has normally developed toes. snout moderately long. on outside of tibia. limbs slate-black with small buff spots behind and below femur. Head as broad as long. compares different. Paratypes. Nic^foro Maria. Specimens Examined COLOMBIA Huila: Paramo de las Papas. 3. tuberculated skin. 13 hand.

in postaxillary region narrower than greatest width of head. chin with three largeVhite |spots. interorbital diameter V/2 times that of upper eyelid. when limbs are laid along the sides. no tarsal ridge. NATIONAL MUSEUM BULLETIN 2 88 lower. heel reaches axilla. vertical. foot. distinct. no apparent skinfold across the chest. with a few small Color in alcohol. third toe a little shorter than fifth. slightly projecting.. 12. Nostrils lateral. soles of feet . a small oval thumb pad present. Body rather stout.5 mm.. head length from angle of jaw. femur. 15 mm. a skinfold on heel and knee. hand. 12. top of slightly separated. head fairly . Head and body.. the center of the belly with many fine transverse wrinkles. when hind legs are bent at right angles to body. ridges.S. a thick glandular ridge from posterior corner of eye above tympanic area ending above angle of jaw. Toes three-fourths webbed. except inner ones which are completely webbed. Skin of upper parts rough with many pointed tubercles on sides of body. knee and elbow are well separated. and along dorsolateral region to groin. a larger round palmar callus. with faint lateral much shorter than second.5 mm. their distance from end of snout equal to their distance from eye. 15 mm. is smooth. loreal region slightly concave. Tympanum hidden. a slight ventral disk (?) no external vocal sacs in the male. 44. fingers extend beyond groin but not to rear of femur. venter finely tubercular on throat and chest and on lower surface of hind legs. first finger . Slate color above and below. edge of upper eyelids with heavy tubercles. Dimensions. and chest and belly with four more pairs arranged quite regularly paralleling the middle and continuing onto lower surface of femur. its diameter equal to its distance from tip of snout. webbed at base. above shoulder insertion. reaching to base of penultimate phalanx of third. 11 mm.5 mm. — — white spots beginning on tip of snout. flaring outward to the upper lip.. prominent... tip of third toe reaches a point midway between groin and end of body. sides of head slate color except for two small light spots on edge of upper eyelid. fingers and toes with swollen tips scarcely wider than the phalanges. normally developed. head width. Fingers moderately long. Canthus rostralis straight. tibia. adpressed. metacarpal tubercles moderately developed on all except last phalanx of each finger. fourth longer than second. where there is a much larger light spot that extends also onto anterior proximal part of femur. when hind leg is adpressed. 16 mm. heels are When arm is when arm pressed backward. Eye large. continued as short light stripes along interorbital region and as well-separated spots behind eye. slightly greater than interval between nostrils. the disk of which reaches to center of antepenultimate phalanx of fourth a very small round inner metatarsal tubercle and a somewhat larger round outer one.124 U. sacrum and limbs.

Ecuador. which is slightly less in the latter. Bufo spinulosus (Wiegman). 1894. — Cope. p. 1858. p. — Nieden. 1891b. — USNM 118220. 3 (fide Cei. straight. p. Critical measurements of the new subspecies are inclusive type of A. 1902. 117. p. It seems clear that. 274). 16. 1875. loreal region concave. 515. Eye large. 107. Head as broad as long. 1926. pp. Atelopus p. 46. 1963. 600 Peracca. 1926. Chile). p. 5. Description. This form is named in honor of our friend and colleague. p. sloping outwards to the upper lip. — p. p. Whymper. p.— Gorham. Canthus rostralis short. erably beyond the lower.— Boulenger. projecting. their distance from end of snout equal to their distance from eye.— Philippi. 19. pi. 24. 14. p. 1894. p. 107. Despax. pi. Philippi. 40. 8 (no text) . 61. J. 37. fig. Fingers moderate. p. a trace . p. 1934c. 1963c. Dr.— FROGS OF COLOMBIA and palms hands of slate — COCHRAN AND GOIN 125 with a few light spots. 1882a.— Parker. 196. 1958. p. footnote. 1904. 7a. 463. p. Colombia. p. 2 Nieden. p. p. 119. 1902. ebenoides except for the head length. interorbital diameter 1% times that of upper eyelid. 83. Otherwise they are very similar in structure and in color. Orton. 1849. laevis. 316 (type locality. Panama. 1926.— Fowler. —Jimfnez de Atelopus — Nieden. 6.— Rivero. an adult male from Guachacal. 83. Phryniscus laevia. Giinther. 417. 1926. its diameter a little greater than its distance from tip of snout. p. p. 1913. p. laevis fig. Tongue onethird as wide as mouth opening. pi. equal to interval between nostrils. 2. 1954. 1875. 1876. 9163c. 1859b. p. p. in the position of the nostril. Neither has a visible tympanum. p. Phrynidium laeve 1868.— Rivero. p. 1876. 1926. sloping backwards to border of mouth in profile. In Meyen. 375. 1882c. p. 43. 83. 83. Phryniscus ignescens Cornalia." Ecuador). 1901. — Cope.— Nieden. these two forms are closely related.— Griffiths.— Noble. 1893. 22 39. oval. p. although separated by the Eastern Cordilleras. — Remarks. 1902. 108. in 1867. 133. — Boulenger. 1922.— Ruthven. figs. 7. Nostrils lateral. 440. — Boulenger. prominent. 1898a. the tips of the digits light also. p. p. 1868. Phryniscus Quito.— Nieden. 83. p. — — Antelopus laevis. pi.— Werner. p. Marinkelle. 375. 1911. 3. snout acuminate when viewed from above. "in humid places around Latacunga near Quito.— Boettger. 1875. figs. la Espada. 164. A (type localities. 1. its posterior border free and unnotched. The new subspecies from Boyaca differs from the nominate form from Huila mainly in the shape and length of the snout. Jiminez de la Espada. 1938. C. — — 1880. 151. p. 1867. or nearly so. and in the incision of the toe webs. — 1926. ignescens. 268. p. the upper jaw extending consid- Narifio. p. 50. of those of the Atelopus ignescens (Cornalia) Plate 22d-f 1834. 139.

either. a heavy skinfold on heel and knee... but separation would not be advisable on that character alone. heels are narrowly separated. Soon after preserving. top of forearm. a little darker on eyeabove canthus. When arm is adpressed. 11. the edge of the mouth pale buff. most having the toes one-half webbed. 31. In Colombia the webs extend from one-third the toe length to the tips. when hind legs are bent at right angles to body. a very present. no ventral disk.S.. head width. but forearm much enlarged. femur. 10 mm. knee and elbow touch. third toe equal to fifth. 9. conical and pointed. Considerable differences in toe webbing are found in both countries. first first finger and second much fingers only. when limbs are laid along the sides. Toes one-half webbed. Ecuador. some of the granules. with an orange Color in alcohol.. belly dirty wax yellow. — hand. a small but distinct oval inner metatarsal tubercle and a smaller round outer one. the tips of the digits and the palmar and plantar tubercles cream color. sides of body. 12 mm. no tarsal ridge. Chimborazo. —A comparison measurements of 36 specimens from Ecuador with a like number from Colombia indicated no significant differences.126 of NATIONAL MUSEUM BULLETIN 288 U. in postaxillary region narrower than greatest width of head. immaculate. crossing posterior part of belly. subarticular tubercles of toes small and irregular. USNM 103242 from Mt.5 mm. Head and body. tip of third finger reaches to end of body. Some specimens of this lot are almost uniform dull peagreen below.. venter smooth. head length. throat maize yellow. but all with lateral shorter than second. The Colombian frogs had slightly wider heads. a few of them double. Skin thick. both reaching to center of antepenultimate phalanx of fourth. web between ridges. but smooth on head and center of back and upper surfaces. 11. buff femoral pattern. In Ecuador the toes are Remarks. Dimensions.5 mm. and on upper limb surfaces. — Dorsum Vandyke brown.5 mm.5 mm. others are sulphur yellow below. side of head clay color. no skinfold across the chest. 9 mm. lids. tibia. metacarpal tubercles well developed. when hind leg is adpressed. fourth longer than and reaching halfway on penultimate phalanx of third. palms of hands and soles of feet wood brown. ending above shoulder. of critical . venter buff anteriorly to olive buff on lower limb surfaces. a short swollen glandular ridge from posterior corner of eye. small thumb pad Body moderately stout. especially those on forearm. and paler along sides of back. heel reaches to front of shoulder. but without any distinct markings. foot. was blackish olive. no external vocal sacs in the male. a large round palmar callus. an orange chrome patch on lower proximal femur. and back of femur coarsely granular.. side of body pale clay color. lightening towards the belly.

its posterior border free and unnotched. interorbital diameter nearly twice that of upper eyelid. Volcan Galeras. 1899.— Rivero. approximately equal to interval between nostrils.. 107.. Fingers moderate in length. Specimens Examined COLOMBIA Cauca: Near Moscopan. the upper jaw extending well beyond the lower. The leg is short in all.000m. Alto von Sibat6. as the heel reaches no further than between shoulder and eye. Patia River side. Encano. fourth longer than either and reaching to base of penultimate phalanx of third. first finger shorter than second.900 ft.800 ft. its diameter equal to its distance from nostril. — 1923. USNM 144927-30. 1926. p. slightly rounded at tip when viewed from above. projecting. Atelopus subornatus Werner.000 m. Eye large. 95177. 146468. a small. one-third as wide as mouth opening. 2.300 m. MCZ 33241.800 m. USNM Guachacal.— Rivero.500 ANSP 25679-705. Mount Chimborazo. scarcely sloping out to the nearly vertical upper lip. 4. 54305. Nostrils lateral. loreal region concave and vertical. — USNM 1963c. 54307-8. ANSP 18282-7. webbed at the base. 83. the greatest frequency in both countries falling at the front of the shoulder. Description. ANSP 25672. 1963c. Ricaurte. C6rdoba: Catival. MCZ USNM CNHM USNM USNM MCZ USNM Atelopus varius subornatus Werner Plate 20d-f 1899. above Pasto. 475 (type localities. the nasal openings very small...800 m. 141991. 10. ECUADOR: Guanote. p. p. their distance from end of snout three-quarter their distance from eye. 3001-3. Tuquerres. Hacienda Gorzon to foot of Mount Pichincha.— Dunn.500 m. Pasto. 118723-4. Pacific side. 18139-54. Cumbal.. CNHM 54304. Cundinamarca Colombia). 107. Machachi. 83. straight. CNHM ft. metacarpal tubercles poorly developed. Patia Riverside. and often only to the axilla.. CNHM 61182-3. Quito. most between one-half and three-fourths webbed.FROGS OF COLOMBIA — COCHRAN AND GOIN 127 a little less than one-half to fully webbed. mountains above Chimbo. ANSP 18236.000-10. Head longer than broad. Nieden. Alto von Sibate" near Bogota. Cundinamarca. Tierra Nigra. 9. 118220.. Cundinamarca. 32340. 475 (type localities. 30 no. 3.... snout straight on the sides. an adult female from Bogota.400 ft. a large round palmar callus. 10. USNM Guitarilla. Toes moderate. 2. ANSP 18136. Pacific side. p.. 2. Rio Sapuyes.— Nieden. 54309. p. 33866-71. the webs prolonged into lateral ridges extending to finger tips. and above Fusagasugd. 1944a. 10. Atelopus flaviventris Werner. upper Rio San Jorge. 2. Colombia. tongue very small. Colombia. near Tuquerres. NariSo: Barbacoa. prominent. . Canthus rostralis sharp.500 ft. 3.230-2. 200 m.. Fusagasugd.. indistinct thumb pad present. 103231-67. p 74. oval. p. truncate and sloping backward to border of mouth in profile.

Skin of fairly developed. . When arm is adpressed. 5. when hind legs are bent upper parts pustular but smooth..128 U. venter smooth. indistinct sepia spots on occiput and near dorsolateral line. Cundinamarca. no inner metatarsal tubercle. 12. but forearm probably at right angles to body. 8 (type locality.5 mm. with small. —Dorsum tawny olive. With so few examples. head length. a round flat gland on side well in front of and above the groin. when hind leg is adpressed. San Isidro. adpressed heel reaches only to the front of the shoulder or to the angle of the jaw. the pustules deepest and most apparent on top of head.5 mm. USNM 95177. and along the sides. limits of variation cannot now be determined. 121.. palms and soles buff. first toe short but distinct and fully webbed. a small round outer one. with a wide sepia spot on outer metatarsal . Specimens Examined COLOMBIA Cundinamarca: Bogota. figs. Magdalena: Santa Marta.. head width.. sacrum and most of upper limb surfaces seal brown indistinctly marbled with a lighter brown venter raw sienna. — Dimensions. no apparent vocal sac in the male. in postaxillary region equal to greatest width of head. This form is in the group having the shortest legs. Atelopus pedimarmoratus Rivero Plate 20g-i 1963. no skinfold across chest. 9 mm. with 3 irregular longitudinal drab spots on center of throat and chest. heel reaches to angle of jaw. NATIONAL MUSEUM BULLETIN 2 88 one-half webbed. 14 mm. 3. 37. ANSP 19749-8. tarsal ridge. third toe shorter than fifth. foot. heels touch.5 mm. no ventral disks. as its heavy sprinkling — of small Remarks.S. a pronounced skinfold on heel and knee. p. sides of body buff with many small russet spots. moderately developed. sacrum. hand. 13. tibia. a heavy bony ridge from posterior corner of eye. pi.. 9 mm. Head and body. Its hand averages shorter than that of any other Colombian form of Atelopus except spurrelli. Body rather no stout. but instead a dark brown spots.. tip of third finger almost reaches to end of body. a clove brown spot on gland just in front of groin. which reaches just beyond base of antepenultimate phalanx of fourth. and its head width averages less than any except varius elegans. knee and elbow touch. can thai and upper loreal regions pale sepia. Color in alcohol. 7. lips immaculate tawny ochraceous. when limbs are laid along the side.. 11 mm. Atelopus pedimarmoratus Rivero.800 ft. Colombia). no dark crossbars on upper femur or tibia. femur. region. snout. ending above shoulder. subarticular tubercles of toes single.

Canthus rostralis concave. posterior corner of eyelids to shoulders . oval inner metatarsal tubercle and a smaller round outer one. when limbs are laid when hind legs are bent at adpressed. 28. lower femur burnt umber. heel reaches front of shoulder. and sides of head and body heavily marbled with seal brown..5 mm. a skinfold on heel and knee. fourth longer than either. along the sides. Tympanum Description. 9. interorbital diameter a little greater than that of upper eyelid. the disk of which reaches to center of antepenultimate phalanx of fourth. but thumb swollen and forearm greatly enlarged. surfaces of upper limbs. and about the same size as that of third finger. belly burnt umber with a few large buff spots.5 mm. Dimensions. then distance from end of snout equal to their distance from eye. Body moderately elongate. 7. its greatest diameter apparently one-fifth that of from eye by an interval equal to its own diameter. lower limb surfaces seal brown Color in alcohol. Eye large. its posterior border free and unnotched. Nostrils lateral. tibia. no external vocal sacs in the male. disk of fourth toe nearly covering the tympanum. a narrow. Fingers moderately long. venter finely pustular. an adult male from San Cundinamarca. — Dorsum fawn with an irregular seal brown and with other dark irregular markings on the sacrum. vertical. X-mark from color. descending to the slightly flaring upper lip. chin and throat cinnamon to russet. which nearly covers the tympanum. heels slightly overalp.5 mm. the upper jaw extending well beyond the lower. rounded. in postaxillary region equal to greatest width of head. head width. scarcely projecting. third toe slightly shorter than fifth. oval. head length.. tip of third finger just reaches to end of body. reaching to base of disk of third. a sharp glandular ridge from posterior corner of eye above tympanum. 8. heavily pustular on back. the distal ones rather weak. proximal metacarpal tubercles moderately developed. a weak skinfold across the chest. with weak lateral ridges. its diameter 1% times its distance from tip of snout.5 mm. 11 mm. separated is when arm is adpressed.. flat palmar callus. sloping backwards in profile. oval thumb pad present. no ventral disk. rounded when viewed from above. foot. first finger shorter than second. right angles to body. knee and elbow touch. 9 mm. a long. — hand. a large. sharp tarsal ridge. Toes one-half webbed. Tongue half as wide as mouth opening. Head and body. sides and upper limb from surfaces.. ending above angle of jaw.. Skin of upper parts smooth on head. When hind leg eye. snout long. Colombia. ending just before the heel. equal to interval between nostrils. femur. Head as broad as long.. Isidro. loreal region concave. a rather flat. rather indistinct.5 mm.FROGS OF COLOMBIA — COCHRAN AND GOIN 129 —CNHM 81872 (holotype). elongate. prominent. free. 10. with small transverse ridges chin to chest.

a weak. longer than broad. vertical above. Description. either. and fifth one-half webbed. similarly dark. Macizo Central Colombiana. Tympanum hidden. in the other. Eye large. ending near heel. a flat round palmar callus. fourth. Fingers rather short. Colombia. tongue very wide as mouth opening. 137531. "between Huila and Cauca. inner ones fully webbed. Atelopus nicefori Rivero. Two rather dried-up specimens (USNM 137531. rounded inner metatarsal tubercle and a slightly stronger round outer one. fourth and fifth toes considerably reduced in size." Paramo de 144936. and the sides of the body are As in so many other species known from but a few examples. the upper jaw extending well beyond the lower. 115. — CNHM Antioquia. and flaring slightly to the upper lip. Toes moderately long. Can thus rostralis sharp and straight. third. the venter is black with some large white spots. 1963c. prominent. its and unnotched. more material is needed in order to gauge variability. Nostrils lateral. a trifle greater than interval between nostrils. third. p. posterior femur fawn color above to dull burnt umber below. NATIONAL MUSEUM BULLETIN 288 with large buff spots. palms of hands and soles of feet wood brown. Caicedo. interorbitul diameter lj^ times that of upper eyelid. fourth midway on penultimate phalanx of finger reaching but not enlarged. Antioquia. longer than free. little 2. third toe much shorter than fifth. 69748 Head a pi. a sharp. only the basal metacarpal tubercles well developed. thin tarsal ridge. about one-fourth as posterior border free lateral ridges. the disks and lower surfaces of toes buff. 144936) tentatively have been identified with this species. (holotype). Remarks. figs. Specimens Examined COLOMBIA Cundinamarca: San Isidro. USNM Catjca: Popayan. subacuminate when viewed from above. its diameter a little greater than its distance from tip of snout. the tip of which reaches to base of antepenultimate phalanx of fourth. it is light with black spots. (Both identified las Papas. first much shorter than second. the back has essentially the same — dark median chevron as in the type. with elongate. In one (USNM 137531).S. USNM tentatively. Colombia). narrow. loreal region concave. snout long.130 U. slanting backwards in profile. In each. disks bulbous taking the place of a . a distinct skinfold on heel and knee. CNHM 81872 (holotype).) Atelopus nicefori Rivero Plate 21g-i 1963. Caicedo. others weak. distinctly projecting. their distance from end of snout equal to their distance from eye. two or three small tubercles thumb pad. 4-7 (type a male from locality.

6977 (paratype). anterior. with a suffusion of wood brown at base of two outer toes. apparently forking and continuing diagonally nearly to the sacrum venter buff. A male paratype (CNHM 81875. from the C amino Sibate-Aguadita) has larger and heavier tubercles and a more distinct marbling on the back. [Choco. head length. 1 (type locality. and posterior limb surfaces burnt umber with slightly darker tips on the small granules where they occur. in postaxillary region a When greatest width of head.5 mm. 7 mm. Gorham.800 m. Atelopus spurrelli certus Barbour. but first finger swollen and elongate. knee and elbow overlap when hind legs are bent at right angles to body.. curved. 12 (type locality. Condoto. sacrum. Specimens Examined COLOMBIA CNHM Antioquia: Caicedo. venter appearing granular because of the crossing of a great number of fine longitudinal and transverse folds..5 mm. Mt. Rivero. no ventral disk. p. no external vocal sacs in the male. near Pena Lisa. 6 mm.. 1923c. pi. The female topoparatype (CNHM 69747) has a throat patch of small dark spots that are better defined than in the male. ..FROGS OF COLOMBIA Body moderately — COCHRAN AND GOIN 131 little less than the hind leg is adpressed. 813. — 1923. Skin of upper parts pustular.. 9 mm. fig. 107. a heavy glandular ridge from posterior corner of eye above tympanum. and the dark-tipped pustules are heavier and more noticeable on the sides of the head and body. Atelopus spurrelli Boulenger. p. — Remarks. ending in a short downward bend above the shoulder. head width. palms of hands and soles of feet drab gray. 25 mm. p.. the pustules mixed with small tubercles on sides. in adpressed arm. Province not specified: Camino Sibate-Aguadita. 25. p. tibia. CNHM Atelopus spurrelli Boulenger Plate 19a-c 1914. 7. and limbs. I. Head and body. — — . side of head drab. femur. third finger reaches only to end of sacrum and does not extend behind body. heel reaches posterior corner of eye. Marx. immaculate but with a russet tinge posteriorly. the throat with a large diffuse V-shaped wood brown patch. 10 mm.] Colombia). . — Dimensions. an indistinct darker Y-shaped mark from posterior corners of eyelids down the center of the back. 8. the belly and lower limb surfaces immaculate. foot. 426. 1963c. 69748 (holotype). superior. a faint skinfold across the chest. a definite dark X-mark on the head. heels touch. 1963. 1958. when limbs are laid along the sides. —Dorsum burnt umber with suffusions of russet on the head.. Sapo. Color in alcohol. eastern Panama). 81875 (paratype). hand. 1. p.

subarticular tubercles of foot small. with small wood brown spots on are laid along the sides. penultimate phalanx of third. NATIONAL MUSEUM BULLETIN 2 88 — USNM Description. 110958. forking to anterior eyelid. Dorsum wood brown.132 U.5 mm. its tip reaching beyond base of eyelid. this light stripe widened above tympanum and containing two small dark spots. foot. and continuing as an irregular dorsolateral stripe from posterior eyelid to groin. penultimate phalanx of third toe reaches of body. no skinfold across the chest. its posterior border free and unnotched. tibia. When arm to end is adpressed. Canthus rostralis rounded and slightly concave. a small round metatarsal tubercle and an outer one of about the same size and shape. a short bony ridge from posterior corner of eye above tympanic area. heel reaches nostril.. and to interval between nostrils. Dimensions.. descending to the Eye upper prominent. when hind legs are bent at right angles to body. Head as long as broad. the first and second reduced in size and fully webbed. slightly projecting..5 mm. free. no tarsal ridge. Fingers moderate. postanal region with a dark brown chevron. first finger not well developed. 23. vertical. without lateral ridges. large. Head and body. with a pale ecru drab median line beginning at tip of snout. its diameter equal to its distance from tip of snout. fourth longer than second. the upper jaw extending well beyond the lower. top of femur with a wide diagonal umber bar. a weak palmar callus. the tip of which reaches beyond base of antepenultimate phalanx of fourth.S. 6. weakly developed. 11. an adiilt from Puerto Utria. head width. and elbow overlap. Choc6. their distance from end of snout about one-third their distance from eye. heels overlap.. anterior femur drab. femur. 12. venter cream buff. no external vocal sacs in the male. single. widening below. the pale ecru drab area surrounding it with two round dark spots. with a sharp demarcation with the paler upper surface. Body very slender. snout bluntly rounded when viewed from above. venter also pustular. the pustules a little larger on posterior belly and below venter. 7 mm. hand. oval. interorbital diameter approximately equal to that of upper lip. 8.5 mm. Nostrils lateral. 7 mm. a very indistinct thumb pad present. each chevron with a diagonal row of dark spots up its center. in postaxillary region narrower than greatest width of head. loreal region flat. Toes moderate.5 mm. no ventral disk. Tympanum not visible. metacarpal tubercles weakly developed.. one-half webbed. and farther back with two rather wide light chevrons connecting the two stripes.. tongue half as wide as mouth opening.5 mm. Color in alcohol. Skin of upper parts minutely pustular. Colombia. flat and slanting backwards in profile. when limbs when hind leg is adpressed. lead length. knee — — . third toe shorter than fifth. a slight skinfold on heel and knee. elongate.

5. both Panama and Colombia. varius bibroni is considered in the discussion under the latter. Puerto Utria. 155. varius elegans and A. 81. 1882a. MLS 172. The light lines on its back are finer and there are fewer spots on its belly. 6. Ecuador. spurrelli certus from nominate — the breaking up the dark dorsal areas of spots. 5. spurrelli of finer spotting of the belly in males. v. p. the and the nearly immaculate belly in females of certus — are subject to much variation in examples from A.5.FROGS OF COLOMBIA and elongate —COCHRAN AND GOIN 133 running longitudinally on belly. head length. olive spots sepia lateral line beginning at tip of snout — all three forms. 19. glyphus in this characteristic. a and running along the canthus. to retain the name spurrelli for all of them. The probable relationship of this species to A. head width. although it is approached by A.). 1926. 1891b. The above-described example is nearly identical with Boulenger's figure of his type specimen from Pefia Lisa. 2.5. 1882c. enlarged on top of head to the dark spots bordering the median light line of snout. tibia. a large brown patch on outer half of tarsus and similar but paler markings on lower surfaces of tibia and femur. p. The adpressed heel reaches between the anterior corner of the eye and the nostril in all except one example. 9. p.— Nieden. 9. it seems best Specimens Examined COLOMBIA Caldas: Manzanares. in which it reaches only to the center of the eye. Dimensions (in mm.5. Solono. Chocd. which was only a half-millimeter longer. The femur averages a greater proportional length than in any other Colombian atelopid. Choc6: El Valle. USNM USNM 110958. The characters by which Barbour separated his subspecies A. The tibia is also very long in throat. hand. text fig. from Manzanares. For the present. Tanti. 7. 6. USNM 151280-9. Ten specimens from El Valle. 134. Caldas) are as follows: head and body length. Phryniscus elegans Boulenger. agree well with the first two mentioned. otherwise it closely resembles the Puerto Utria example. palms of hands and soles of feet cream buff. femur.) of a young frog (MLS 172. except that two of them have nearly immaculate bellies. Choc6. Atelopus varius elegans (Boulenger) Plate 19d-f 1882.000 ft. Bahia 144933-5.5. continuing and widening greatly behind eye to groin. foot. below and in front of eye. upper lip cream buff with two wood brown spots on its margin. p. Remarks. with wood brown spots across penultimate phalanges of the three outer toes and fingers. 464 (type locality. 337-262—70 10 . therefore.

pp.— Griffiths. 1926.5 12. 16 mm. p. 32. a heavy skinfold on heel and knee. p. one-half webbed. knee and elbow overlap. curved slightly. 83. which reaches hallway along the antepenultimate phalanx of fourth. 553. with lateral ridges. 81. no skinfold across the chest. Body slender. foot. — 8. Skin of upper parts thin. its diameter equal to its distance from tip of snout.— Parker. 24.5 mm.third their slightly projecting. vertical. Head and body. p.— Rivero. 1894.— Gorham. Can thus rostralis distinct.S. Atelopus elegans. Toes long. 1963. Boulenger. p. Description. which is also rather short.. head length.— Dunn. 1926. 1926. sloping backwards to mouth in profile. but forearm greatly thickened near elbow. third toe shorter than fifth. in postaxillary region narrower than greatest width of head. 518. 8 mm. Colombia. from end of snout one.. its posterior border free and unnotched. USNM . a faint. when limbs are laid along the sides. p. base of second finger reaches to end of body. p. no subarticular tubercles on foot. 9 mm. a small.—Parker. descending to the vertical upper Eye lip border.— Nieden.— Nieden. tibia. the upper jaw extending well beyond the lower. 119. tongue one-third as wide as mouth opening. so that arm cannot be straightened. p. 1954. a few granulations on sides and sacral region. 553.. Colombia. a larger round palmar callus. no ventral disk. 1898a. 47. p. prominent. 52401 is badly has been chosen for the color description. p. Nostrils lateral. NATIONAL MUSEUM BULLETIN 2 88 — — Peracca. Colombia) . p.5 mm.134 1894. since Color in alcohol. 1904. about one-third to onefourth of the total length.. heel reaches a point between eye and nostril. when hind leg is adpressed. an adult female fiom Gorgona Island. 375. weak thumb pad present. first finger not well developed and much shorter than second. halfway on penultimate phalanx of third. mm. interorbital diameter 1% times that of upper eyelid. elongate. 102 (type locality. p. 1905. 22. a weak glandular ridge from posterior corner of eye ending above shoulder. webbed at base. oval inner metatarsal tubercle and a more distinct round outer one.5 mm. Atelopus gracilis Barbour.. heels overlap. head width. no tarsal ridge.. (ANSP 23177 from Gorgona Island. oval. when hind legs are bent at right angles to body. snout rounded at tip when viewed from above. Fingers long. 40. 107. — Dimensions. their distance distance from eye. no external vocal sacs in the male. 1926. 1963c. anteriorly nearly smooth. Gorgona Island. U. reaching nearly large. fourth finger normal. equal to interval between nostrils. venter anteriorly with many minute transverse wrinkles on throat and chest. hand. Head a little longer than broad. metacarpal tubercles not developed. When arm is adpressed. 1944c. femur. 14. loreal region concave. — USNM 52401. granulate on posterior belly and lower femur.

. this light stripe bordered above and below by clove brown stripes. the upper jaw extending well beyond the lower. third and fourth distinctly webbed Tambo. ECUADOR: Solidero. USNM 118219. flaring very slightly to the upper lip. Giiisito.— CNHM Tambo. the upper lip white. palms and soles immaculate cream-buff. 50973-82. ANSP 23176-7. Eye large. Fingers moderately long. vertical. the second. above tympanic area and merging with a white inguinal spot. The venter is immaculate in all specimens.) and somewhat broken dark brown stripe on either upper surface of legs and arms fawn color with a network of dark irregular lines not forming real crossbars. MLS 173. a less distinct side of midline from snout to anus. groin. its posterior border free and unnotched. snout broadly obtuse when viewed from above. this tone sharply divided from the loreal color. — Specimens Examined COLOMBIA Cauca: Gorgona Island. pi. Tympanum hidden. 145134-230. as well as the more or less irregular dark broken stripes on either side of the midline. elongate. 145132. MCZ 52401 (cotype of Atelopus gracilis Barbour). slightly rounded. as practically all examples at hand show the white dorsolateral stripe edged with dark brown stripes above and below. an adult female from El Cauca. p. anterior and posterior femur buff. Head as broad as long. 1. 800 m.FROGS OF COLOMBIA —COCHRAN AND GOIN 135 Dorsum cinnamon. Canthus rostralis straight. El Giiisito. Nostrils lateral. 8 (type locality. Cauca. described above. 49802-11. side of head clove brown in loreal region. its diameter equal to its distance from tip of snout. 112. there is great uniformity in the pattern elements. Atelopus longibrachius Rivero Plate 22a-c 1963.. covering nearly half the side between axilla and faded. oval. Atelopus longibrachius Rivero. 7605. AMNH 2615. upper eyelid. Variations in the critical measurements are normal. Colombia. with lateral ridges. Colombia. their distance from end of snout one-half their distance from eye. considerably greater than interval between notrils. Description. prominent. the lower brown stripe wide. unspotted. interorbital diameter nearly twice that of upper eyelid. however. fig. Pacific side). 54283 (holotype). venter immaculate cream-buff. with a white (yellow) dorsolateral stripe beginning at snout tip. tongue one-third as wide as mouth opening. loreal region weakly concave. continuing on canthus. sloping backwards in profile. moderately projecting. Some individuals are a little darker or lighter than ANSP 23177. Remarks.

broken dorsolateral stripe beginning at tip of snout. Toes moderately long. venter weakly granular posteriorly. The adpressed in heel reaches to the center or anterior corner of the eye most examples. suggestions of a skinfold across the chest. 12 mm. rounded inner metatarsal tubercle and a slightly smaller round outer one. heels touch. and body. its disk reaching to base of penultimate phalanx of fourth. when hind legs are bent at right angles to body.136 U. chin and anterior half of belly immaculate. fingers extend behind body by at least the two terminal phalanges of third finger.. — Dorsum. ending midway to heel. rest of side of head olive buff.5 mm. 10 mm. a skinfold on heel and knee. in postaxillary region equal to greatest width of head. mm. and a few small irregular sepia ocellated spots on its posterior half. a larger round palmar callus. —The large series uniformity in color pattern. Skin of upper parts smooth. fifth toe longer than third.. first NATIONAL MUSEUM BULLETIN 288 and second completely webbed. the throat. continuing below canthus. knee and elbow overlap. heel reaches center of when arm is adpressed..5 mm. and fifth almost completely webbed. no ventral disk. the others completely webbed. immaculate.. —Head Dimensions. fourth. venter light clay color. pustular eye. first finger reduced. reaching halfway on penultimate phalanx of third. head width. concentrated into an irregular. head length. much shorter than second. belly. When hind leg is adpressed. metarcarpal tubercles present. tibia. a small flat.S. third. fourth longer than second. no external vocal sacs or thickened forearm in the male. femur. lateral stripe. rather weak glandular ridge from posterior corner of eye above tympanic area.. 10 hand.5 millimeters in total length. irregular sepia ocelli on anterior and posterior limb surfaces. Remarks. Body elongate. and upper limb surfaces brown with a coarse but regular sepia vermiculation which is wood Color in alcohol. ending above shoulder insertion. fingers and toes with small swollen disks scarcely under the phalanges. when limbs are laid along the sides. but a smooth brown callosity on inner surface of first finger. 18 mm. foot. a weak tarsal ridge. at base. 14. although it may come only to the angle of the jaw . with many fine transverse and logitudinal wrinkles anteriorly. palms of hands and soles of feet raw sienna. 38 mm. the anterior and posterior region of the eye sepia. under the lens.. the pustules becoming small granules on sides of head and body and on sacrum. a wide. above tympanic area and along side of body to groin. 16. sides of body. of Many 26 paratypes shows remarkable lack the small dark spots on the and some have a wide wood brown area above the dark dorsoThe largest example is 43. a small oval thumb pad present.

fourth longer than either and reaching halfway on penultimate phalanx of third. a heavy skinfold on heel and knee. 3. No tarsal ridge. . Body rather elongate. Nostrils lateral. pi. and upper limb surfaces. when hind leg less adpressed. Outer toes one-half webbed. webbed at the base. the upper jaw extending far beyond the lower. Rio San Juan. Rivero. Boulenger. and — COCHRAN between eye and to a point skin at the edge of the upper eyelid is AND GOIN nostril in still fewer. p. 147505. and with the tip projecting in profile. Head a little longer than broad. is with numerous small conical tubercles on sides of body. when hind legs are bent at right angles to body. p. single. 375. Antioquia. Specimens Examined COLOMBIA El Tambo. a small round thumb pad present. Atelopus varius bibroni (O. when limbs are laid along the sides. a small oval inner metatarsal tubercle and a smaller round outer one. 1926. When arm is adpressed. 1963. Pacific side. Cope. venter finely granular on throat and chest. 14. 800 m. interorbital diameter 1% times that of upper eyelid and a trifle wider than interval between nostrils. New Granada [Darien region of Colombia and Panama]) 1858. a large round palmar callus. p. fig. Schmidt) Plate 21d-f 1857. 256. Eye large. 1926. its tip reaching almost to penultimate phalanx of fourth. tip of third finger reaches a point between groin and end of body. prominent. developed than those of hand. 84. 1963c. 84. Schmidt. obovate. p.[telopus] bibronii. . first toe distinct but entirely webbed. Tongue slender. with very wide lateral ridges. p. 1867. subarticular tubercles of foot small. slightly projecting. knee and elbow are separated. p. Can thus rostralis sharp. Cauca: CNHM CNHM 54283 (holotype). sacrum. 137 The only slightly thickened. p. 196. 400 m. — — — USNM Description. 25 (type locality. snout obtusely pointed when viewed from above. continuous with the upper lip. Choc6: Playa de Oro.. heels touch. 1894. a female from Medellm..— Nieden.— FROGS OF COLOMBIA in a few. Giiisito. Hylaemorphus bibronii O. its posterior border free and unnotched. in postaxillary region narrower than greatest width of head. metacarpal tubercles well developed. 108. 54284-304 (paratypes). Colombia. A.— Nieden. its diameter equal to its distance from tip of snout. their distance from end of snout two-thirds their distance from eye. 54276-82 (paratypes). Phrynidium bibronii. loreal region slightly concave and vertical. Skin of upper parts heavily glandular. one-third as wide as mouth opening. fifth toe a little longer than third. 1867. heel reaches to angle of jaw. first finger much shorter than second. Fingers moderately long.

with a lighter area in front of eye extending to border of upper lip and a more distinct clay colored spot at commissure of jaws. 13 mm. with the webs and tubercles light clay color. 11. 33 mm. and the remainder have large or small light dorsal spots. 8. femur. USNM 147505. Cauca: El Tambo. CNHM 61755 (juv. MLS 312-3. Remarks. spurrelli in body proportions as well as Both have the wide dark brown lateral stripe. in- in color pattern. Four of the other examples from Antioquia are immaculate vandyke brown above. a short. no ventral disk. tibia. while the tip of its snout is less pointed than others of similar size from Antioquia. . head length.. foot. Antioquia. 2 88 coarsely granular on belly and lower femur. Antioquia: Andes. but forearms thickened near elbow. do not warrant its separation as a subspecies. no crossbars on legs. no external vocal sacs in the male. head — width. with a serrate border where this color meets the ventral tone.. mm.). like the described specimen. These small differences. ANSP 25365-8. Pereira. usually concentrated on the snout and parietal region. but in v. —There is a definite suggestion of relationship between Atelopus varius bibroni and A.5 mm. Colombian Specimens Examined USNM 144931-2 (juv. AMNH 3208. Medellin. Head and body. No skinfold across the chest. hand.5 8. 1345-6. bony ridge from posterior corner of eye above tympanic area. bibroni the back (between these stripes) has no pale chevrons outlining more or less regular dark patches. sides of body vandyke brown.. Narino: Ricuarte.). 1377-8. lower surfaces of arms and legs pale sepia except for proximal area of femur including postanal region.138 NATIONAL MUSEUM BULLETIN U. 13537-8. chin and throat sepia. Amazonas: Rio Apaporis.. — Variation. 25671 (tentatively identified). Dimensions. side of head dull vandyke brown.5 mm. ANSP 25369 (tentatively identified). and with a sharp division between the two colors across the lower proximal part of femur. which is clay color like the belly.. MLS 170 from Cundinamarca has a slightly smoother skin. ANSP 25362. : Caldas: Laguneta. Dorsum immaculate Vandyke brown in this specimen. 10 mm. 14 mm. however. ending just above angle of jaw. there are irregular elongate dark spots arranged haphazardly in three rows. stead. 39088. MLS 171. Cundinamarca: MLS 170. BoLfvAR(?) Probably upper Sinu River. belly light clay color. but a darker brown color around the base of each tubercle. the lighter examples of A. heavy. — Color in alcohol.. with a more or less apparent light dorsolateral line and with faint traces of dark and light crossbars on the legs. palms of hands and soles of feet sepia. Quindfo. or the back may be uniformly dark as in the described specimen. AMNH 1340. AMNH 14149.S.

5. when limbs are laid along the sides. a blunt oval iimer metatarsal tubercle and a smaller round outer one. tongue approximately two-fifths as wide as mouth opening. When arm is adpressed. where it ends in a swollen gland. concave. its length about one-third of total length.5 mm. 518. Body fairly stout. palms and soles slightly lighter gray.— Rivero. 1922. p.. 15. pi. 17 mm. 15. head length. tip of third finger reaches to end of body. its posterior border free and unnotched. 19 mm. Head as long as broad. head width. subarticular tubercles mostly double. first finger shorter than second.] Colombia. 1. 107.. an adult male from Don Description. Nostrils lateral. but forearm greatly enlarged near elbow. no external vocal sacs in the male. Alelopus carrikeri Ruthven. Dimensions.5 mm. p. no tarsal ridge.. Skin of upper parts thick but smooth. loreal region vertical. 18. Outer toes one-half webbed. ventor smooth. 1963. stalked. extending on shoulder and to forearm and elbow. 1963c. in postaxillary region a little narrower than greatest width of head. No skinfold across the chest.5 mm. free. 14 mm. Dorsum uniform blackish slate. p.. knee and elbow are separated. Magdalena. Dunn. — — . venter uniform slate gray excepting for a pale round spot on each side of chin below mouth commissure. 1944c. rounded in profile. scarcely projecting. the upper jaw extending well beyond the lower. prominent. not strongly developed. third toe a little shorter than fifth. short. 1 (type locality. entirely webbed. a small oval thumb pad present.— Gorham. tibia. Color in alcohol. first toe distinct.. p. thick. heels are separated. a low glandular swelling from tip of snout along canthus to posterior tympanic area. snout rather acuminate when viewed from above.. foot. no ventral disk. Head and body. its lower part flaring very slightly to the upper lip.5 mm. Eye large. its diameter four-fifths its distance from tip of snout. Fingers thick. —MCZ 4100 (paratype). heel reaches slightly beyond axilla. Diego. Santa Marta Mountains. Paramo de Macostama. hand. Colombia. fourth longer than either and reaching halfway on penultimate phalanx of third. a patch of small spicules behind lower jaw. femur. interorbital diameter 1% times that of upper eyelid and a trifle greater than interval between nostrils. Can thus rostralis rounded but strongly developed. 24. their distance from end of snout equal to their distance from eye. when hind legs are bent at right angles to the body. fairly long. [Magdalena.— FROGS OF COLOMBIA —COCHRAN AND GOIN 139 Atelopus carrikeri Ruthven Plate 22 g-j 1916. the tip of which reaches halfway on antepenultimate phalanx of fourth. 51. a large round palmar callus. a heavy inner skinfold on heel and knee. p. metacarpal tubercles absent. when hind leg is adpressed. with lateral ridges.

Breder. Fingers long. loreal lateral.000 m. reaching penultimate phalanx of second. 1946. The specimens examined are dark brown to blackish 123560 (see Plate 22j) which has a dorsal above except pattern of striking large black and white (yellow in life) spots. Dari£n. with vertical phalanges of fourth free. MCZ 4100. weakly developed. and 145744glaciers. near head of Rio Lim6n.140 U. webbed at base. p. NATIONAL MUSEUM BULLETIN 288 — Remarks. metacarpal tubercles not developed. an adult male from the of Rio Lim6n. p. with faint lateral ridges. Rio Guatapuri. Darien Province. Magdalena: AMNH USNM USNM USNM Atelopus varius glyphus Dunn Plate 21a-c 1931. squarely truncate when viewed from above. 50965-7. even considering the fact that the series examined consists of only 11 examples.800 m. In the critical measurements. — — USNM 50230 (holotype). its disk reaching a little distance along antepenultimate phalanx of fourth. first finger very small but distinct.third as wide as mouth opening. In all other respects 123560 resembles the other specimens from USNM slate USNM Magdalena. variation is not great. their distance their distance from eye.S. in postaxillary region a little narrower than greatest width of last three small . Eye large. its posterior border free and unnotched. 409.. 4. one. Pirri Range. 123560-2. fourth longer than second. 4. interorbital diameter 1% times that of upper eye lid. slightly projecting. straight and nearly vertical in profile. first and second entirely webbed. Uscategni. reaching halfway on antepenultimate phalanx of third. a blunt oval inner metatarsal tubercle and a smaller round outer one. third a trifle longer than fifth. a distinct skinfold on heel and knee. thumb pad and palmar callus very faint.900 ft. merging with rostralis the nearly upper lip. Tongue small. its diameter three-fourths its distance from tip of snout. a little greater than interval between nostrils. oval. the upper jaw extending considerably Description. 118725-6. Specimens Examined COLOMBIA Don Diego. Body elongate. Canthus region vertical concave. Mamancanaca Valley. prominent. first toe small but distinct. Sierra Nevada de Santa Marta. 10. Head a little longer than broad. 5. third to fifth one-half webbed. Pirri Range near the head beyond the lower. their tips swollen into small disks. toes scarcely swollen into disks. snout moderate. Panama).. Paramo de Surubaka. Toes moderate. Sierra Nevada de Santa Marta. Nostrils from end of snout one-fifth distinct. subarticular tubercles of toes single. Panama. no true tarsal ridge. 396 (type locality.. Atelopus varius glyphus Dunn.

a bony ridge from posterior corner of eye ending above shoulder. tibia 19. Dorsum cinnamon. 1875. 53964 (paratype). Skin of upper parts smooth. ANSP 24245. 54191-6 (paratypes). Rio Jappe. none of which seem to be very old. USNM 48594-5. Some of the paratypes that are smaller and younger than the type have rather elaborate patterns.. 4. 11. Atelopus longirostris Cope Plate 12d-f 1868. and followed by a large squarish gland just above arm insertion. except for some small transverse glands on sacrum.. a slight skinfold across chest. 18. Dimensions. knee and elbow touch. Atelopus longirostris Cope. immaculate. 2.5 mm. 50168-75. 155. figs. 1-ld. venter light ochre yellow. 140058-9. no external vocal sacs. 17 mm. head width. when limbs are laid along the sides. head length. 50231-48 (paratypes).. dark interorbital patch.— FROGS OF COLOMBIA — COCHRAN AND GOIN 141 head. femur. 54230 (paratypes). Pirri Range near head of Rfo Lim6n. being broken for only a short distance on the lower femoral surface. — .5 mm. canthal and loreal regions dark brown. the ground color of dull sage green is nearly obscured by three very wide clove brown chevrons.5 mm. p. Dari6n Province. upper eyelids and gland above shoulder pale ochre. Valley of Quito. Head and body. hand. Remarks. venter smooth. 44 mm. similar dark crossbands nearly encircle the legs. canthal region. 13. except for weak granules below anus.. heel reaches posterior corner of eye. Usually the toes are about one-half webbed. except that the small brown dots on the belly are more scattered ANSP 25626 lacks ventral spots. The Colombian specimens. Porto Obaldia. foot. heels slightly overlap.5 mm. USNM PANAMA: USNM USNM USNM Chiriquf. the rest of the upper lip olive buff. uniform except for some small Color in alcohol. venter olive buff with a double row of small brown spots along its midline from chin to posterior abdomen. pi. 12. tip of second finger reaches to end of body. Siola. when hind leg is adpressed.. Panama. 116 (type locality.5 mm. Jimfnez de la Espada. upper lip.200 ft. Ecuador). 50226. When arm is adpressed. 50230 (holotype). a few indistinct olive crossbars and spots on legs. near Cana. the point of the anterior one produced into a wide. from Cerro Azul. when hind legs are bent at right angles to body. scattered guanine spots. a faint ventral disk. — — — USNM USNM Specimens Examined COLOMBIA ANSP 25626-31. Cerro Azul.. 2-2c. but the webbing appears to be more deeply incised in some individuals. Cana. p. In 54191. resemble 54191 very closely. Darien. but forearm of male developed for clasping. Choc6: Upper Rfo Jurubida.

spatulate. due to elongated snout. 11. Nostrils lateral. apparently a tarsal ridge. a small oval thumb pad present. NATIONAL MUSEUM BULLETIN 1876. heel probably reaches to center of eye (not possible to acertain in the stiffened specimen) when limbs are laid along the . adhering to vertebrae. p. 2 p.S. the upper jaw extending con- siderably beyond the lower. a skinfold on heel and knee. Orton. — — CNHM Description. p.. Rivero. 43850. snout bluntly pointed when viewed from above. finger disks bulbous but not greatly enlarged. 1901. femur. 14 mm.— Rivero. 1963. descending almost vertically to the upper lip. 107. 600 (type locality.. 25. ending near heel. which are very prominent. p. its posterior border free and unnotched. Head and body. tibia. the outer two entirely webbed.— Gorham. knee and elbow touch.—Werner.. 19.. Fingers rather long. and sharp. slender. 1963c.— Nieden. p. with lateral ridges. Colombia.— 142 U. its length about 1% times its width. an adult male from Cisneros. 108. in postaxillary region equal to greatest width of head. 1901. third and fifth subequal. 19. first and second one-half webbed and quite short. nearly right-angled at tip. p. prominent. head — width. p. separated from eye by an interval equal to its own diameter. a sharp glandular ridge from posterior corner of eye. venter finely pustular. Nieden. 1926. Tympanum not very distinct. Ecuador). Valle. . foot. head length. its greatest diameter one-third that of eye. projecting above tympanum. a flat.. loreal region slightly concave. Eye large. when hind legs are bent at right angles to body. metacarpal tubercles poorly developed. 14. Atelopus longirostris marmorata Werner.. 1963. 1963c. a small oval inner metatarsal tubercle and a larger rounded outer one. small. sides. 375. 88 1894. hand. Skin of upper parts thin. footnote. fourth longer than either and reaching to base of penultimate phalanx of third. no ventral disk. one-half webbed. their tips reaching to base of antetheir distance Can thus from end of snout equal to their distance rostralis straight penultimate phalanx of fourth toe. ending above shoulder. a light skinfold across the chest. Head flat above.5 mm. Toes long. 100. the three outer ones webbed at base. Atelopus l. When hind leg is adpressed. round palmar callus. 81.5 mm. its diameter about seven-eighths its distance from tip of snout. concave and receding in profile. 13 mm. interorbital diameter nearly three times that of upper eyelid. from eye.5 mm. Dimensions.5 mm. 1926. very long.5 mm.[ongirostris] marmoratus. with coarse pustules on chin and throat and wrinkles on belly. heels touch. 600. scarcely projecting. p. pustular under the lens. 81. Body elongate. first finger much shorter than second. 46. a little greater than interval between nostrils. p.— Boulenger. tongue a little less than half as wide as mouth opening.

with a coarse sepia X-mark from the upper eyelids to a point well behind the level of the axillae. 43850-1. rather sharply set off from the paler ventral color. immaculate on belly. its borders irregular on the body. The other specimen from Cisneros. CNHM Specimens Examined COLOMBIA CNHM Valle: Cisneros. a cinnamon line along the back from nuchal region onto the sacrum. swelling of that digit in the male. USNM 144937-40 (tentatively identified). soles of feet and palms of hands deep brown to black on outer toes. Santander: Hacienda Barrancas. The throat and chest have a claycolored outline of a "chevron" consisting of an irregular median line from tip of chin to breast. upper limb surfaces clove brown to black. lightening to vandyke brown below. The faint tarsal fold mentioned by Cope apparently exists in 43851 as a thin ridge of skin. No external vocal sacs apparent in the male. these examples "more or agree" with Cope's original description. cannot be checked. The first inner finger enlarged. but the ridge on this shrunken and dried specimen may be much stronger than the ridge on fresher material. of Rio . a female distended with eggs) is lighter in color than the male (CNHM 43850) being wood brown above and on the sides. side of head clove brown with an elongate cinnamon spot on upper lip in front of eye and a larger round spot below tympanum. Valle (CNHM 43851. 900 m. Velez. with a cin- on on postocular ridge and along side of body to groin.FROGS OF COLOMBIA Color in alcohol.. with a large chevron-shaped russet patch covering most of chin and dying out on chest. p. but first finger is distinct in the female. 100 m. Quebrada La Lechera. lightening to wood brown below inner digits and on their tips. The venter is olive-buff. anterior femur with an irregular ecru drab stripe edged below by a vandyke brown stripe continued from side of body. They are so badly stiffened and mutilated that Cope's statements as to the extension of the fore less and hind limbs.. with other spots in an irregular ( ) -shaped arrangement on each side. eyelid. namon — Dorsum — COCHRAN AND COIN 143 clove brown. Markings on the limbs are clouded and upper stripe beginning at tip of snout. posterior femur with alternating dark and light squarish spots. sides of body clove brown above. Remarks. The more but this is due to the and second fingers in both male and female are much broader and have wider tips than the third and fourth fingers. tributary Op6n. venter buff. backward and forward. — . 112) has remarked. indistinct. As Rivero (1963c. almost black. continuing along can thus. with some cinnamon spots especially prominent on upper arm.

their tips bulbous. Nostrils lateral. first finger much shorter than second. Fingers rather long. Eye large. pi. an finger when adpressed reaching laid along the sides. a weak middorsal ridge.5 mm. when hind legs are bent at right angles to body. third stout.. Skin of upper parts fairly smooth (pustular under lens). to anus where it forks and continues on back of femur. prominent its diameter a trifle less than its distance from tip of snout. strongly projecting. with many small sepia dots on the head tubercles. a male from Magdalena. third and fifth subequal. NATIONAL MUSEUM BULLETIN 2 88 Atelopus walkeri Rivero Plate 20a-c 1963. 10.144 U. with some low tubercles on head and a parotoid swelling behind the eye.. heavy skinfolds on heel and knee. 1963c. descending to the upper lip. before reaching heel. slightly bulbous. 117. figs. heel reaches posterior border of eye. — width. its posterior border free and unnotched. — Dorsum light clay color. femur. interorbital diameter twice that of upper eyelid. metacarpal tubercles well developed. heels are slightly separated. external vocal sac in the male. loreal region concave.. in postaxillary region equal to greatest hind leg is just beyond groin. curved. 12 mm. vertical. venter smooth except for some minute transverse wrinkles on belly. a heavy glandular ridge from posterior corner of eye along parotoid area. head mm. Body fairly width of head. 39 mm. free. fourth longer than either and reaching nearly to base of penultimate phalanx of third. and granules below anus. a weak skinfold across the chest. a heavier lateral sepia stripe from tip of dark dots . flat palmar callus. forming a broken midline from snout tip Color in alcohol.. a small oval thumb pad present. Don —MZUM 48007 (holotype).5 mm. Tongue one-third as wide a mouth opening. head length. p. 3. the upper jaw extending far beyond the lower. hand. when limbs are knee and elbow just touch. tibia. 15. much greater than interval between Tympanum hidden. Colombia. Santa Marta region. ending above shoulder.S. Description. Atelopus walkeri Rivero. the tips with weak lateral ridges. Can thus rostralis distinct. their distance from end of snout nearly as great as their distance from eye.] Colombia). foot. a slight ventral disk. When adpressed. their tips reaching beyond base of antepenultimate phalanx of fourth toe.5 mm. Toes moderately long. a small oval inner metatarsal tubercle and a smaller rounded outer one. Dimensions. a large. 11 Head and body. oval. straight and slanting backwards in profile. a faint chevron above shoulders. Diego. 1-4 (type locality. one-half webbed. ending nostrils. snout broadly acuminate when viewed from above. [Magdalena. a weak tarsal ridge. 15 mm. 15...

seemingly arranged in broken transverse series. dl . the tips of the digits a tions of pale sepia spots tibia little and foot. Roof of skull exostosed. and claw-shaped terminal phalanges. The similarity of A. walkeri. The holotype is the only known example of A. the two colors irregularly divided. e e 1 . lower surfaces of tibia and foot with sepia vermiculations palms of hands and soles of feet olive-buff. along sides to groin. A. concentra- on upper proximal half of femur and across upper half of posterior femur pale buff. No teeth on parasphenoid. walkeri and A. Atelopus walkeri resembles the Venezuelan A. From the four other families of frogs with intercalary cartilages this family may be distinguished as follows: from the Centrolenidae in having the terminal phalanges claw-shaped rather than T-shaped. between the ultimate and penultimate phalanges. cruciger vogli is rougher dorsally and has longer limbs. side of head mottled with pale sepia. A well-developed helmet No well-developed helmet Cerathyla Osteocephalus . 2 . a double condyle on the coccyx. Vocal pouch paired. edge of upper lip white with a darker sepia line just above it. — Remarks. FROGS OF COLOMBIA — COCHRAN AND GOIN 145 of snout along can thus. and the inner and outer metatarsal tubercles are distinct only when the foot is bent. subornatus also was discussed by Rivero. cruciger cruciger rather closely.. c1 . a large sepia postanal patch. to Colombian Genera of Hylidae Pupil horizontal. The only available Bogota example of the latter (USNM 95177) has the third finger extending slightly beyond the end of the body. but this may be due to the preservation. The tubercles below the digits are distinct. and cruciger cruciger has a very distinctive dark dorsal pattern. from the Pseudidae in having the intercalary cartilages disk-like rather than elongate and rod-like. and larger ones on belly. arciferal girdle. and from both the Rhacophoridae and Phrynomeridae in being procoelous and arciferal rather than diplasiocoelous and firmisternal. behind eye. darker. lower half sepia. although oxyrhynchus has larger glandular warts on the flanks and less webbing between the toes. olive buff with small sepia spots on chest. as the adpressed heel reaches the nostril or snout tip and the third finger reaches well beyond the end of the body. As suggested by Rivero. oxyrhynchus and A. short intercalary cartilages Key a1 . 1 ft . Family Hylidae Frogs with procoelous vertebrae.

toe shorter first toe longer than second (except Phyllomedusa Genus Agalychnis Cope 1864. calcarifer). p. Agalychnis Cope. first than second Agalychnis Palpebral membrane not reticulate. b2 NATIONAL MUSEUM BULLETIN U. Lutz. p. The frogs of this genus may be characterized by the following combination of characters: vertical pupil. Nieden. — — . 181 (type species. Hyla callidryas Cope). c2. Northwestern Ecuador). 1931. of a number of longitudinal folds bounded anteriorly by transverse folds. 601. bicolor) b 2. 1957. 52 (type locality. 1950. 61 . 23. Phrynohyas .146 d2 . with a strong . vocal pouch of male median and subgular. l . 1 A a2 No of the species. region and e2. e 1 . B. p. d Sacral diapophyses expanded. 1923. e Males with vocal pouch on the posterior margin of the throat l . — 1963. Vocal pouches with no tendency to be paired. if subgular. Dunn. palpebral membrane not reticulate. eyes red in life in many Key a. 335. d2 . — Generic diagnosis. — 24. Phyllomedusa calcarifer. in most species snout not strikingly pointed and projecting Hyla Sacral diapophyses rounded. made posteriorly Hyloscirtus . Agalychnis calcarifer Boulenger. 415. in P. Smilisca 1 a2 . Tympanum quite distinct Cryptobatrachus Tympanum indistinct or absent Vocal pouches either at angles of jaws tendency to be paired.S. or. c1 . palpebral membrane reticulate. p. Pupil vertical.—Gorham. to Colombian Species of Agalychnis prominent triangular dermal appendage on heel. p. Funkhouser. d Vocal pouches at angles of jaws d2 Vocal pouches not at angles of jaws e2 c2 . A. webbing more or less well developed on hands. p. first toe shorter than second. calcarifer . three phalanges of fourth toe free or nearly free of webs. 2 88 Vocal pouch unpaired Teeth on parasphenoid Gastrotheca Tetraprion Roof . Rio Durango. p. calcarifer. prominent triangular appendage on heel. spurrelli Agalychnis calcarifer Boulenger Plate 23a 1902. of skull not exostosed. A. palpebral membrane reticulate in all except A. Palpebral membrane reticulate (except in A. tongue extensively free behind. snout pointed and pro- jecting Sphaenorhynchus Males with vocal sac single but not bounded anteriorly and posteriorly by transverse folds. 1950. no more than one phalanx of fourth toe entirely free of web.

finger-like projections from the dorsal ground color onto the ventral light color. which is considerably longer than second. a sloping snout. no projecting rudiment of a poll ex. sloping in profile. Eye moderate. From spurrelli it differs in having a prominent triangular dermal appendage on the heel and a glandular fold above the vent. a well-developed tarsal ridge extending from tip of fifth toe along outer margin of foot to heel where it terminates in a prominent triangular dermal appendage. in postextensively —BM fingers. very slightly projecting. the web on fourth toe reaching the middle of the antepenultimate phalanx. In pattern it differs in having no reticulations on the palpebral membrane. in the presence of brown patches on each side and below the vent. third and fifth toes subequal. which more than covers the tympanic area. slightly elongate series. in having blackish. interorbital distance slightly greater than width of upper eyelid. web reaching middle of penultimate phalanx on lateral side of fourth finger. Condoto. about three-fifths the diameter of eye and nearly in contact with it. the upper jaw not projecting beyond lower. Colombia. which is not particularly wide and about equal to distance between nostrils. in having pigment under the fourth finger and under the metatarsal region of the fourth and fifth toes. first toe shorter than second. from Pena Lisa. a little closer to tip of upper lip than to eye. Toes more than one-half webbed. palpebral membrane not reticulate. loreal region somewhat concave and nearly vertical.FROGS OF COLOMBIA Diagnosis. reaching to middle of disk of third. Can thus rostralis slightly defined. and in lacking the clear white. spurrelli by the presence of extensive webbing between the fingers. its dismeter equal to its distance from nostril. Tympanum very distinct. a distinct oval inner and a very small but distinct rounded outer metatarsal tubercle.12. Fingers extensively webbed. and a triangular dermal appendage on the heel. Vomerine teeth in two small. Description. not opposable. a well-developed ulnar ridge extending from tip of fourth finger along outer arm to elbow. Choc6. — COCHRAN —An Agalychnis with the webbed first AND GOIN 147 toe shorter than the second. tongue half as wide as mouth opening. broadly triangular when viewed from above. separated from each other by an interval slightly less than their distance from eye. elongate. This species differs from all other Colombian frogs with vertical pupils except A. scattered spots on the back. disk of fourth toe practically covering the tympanic area. nostrils more lateral than superior. squarish choanae. the outer ends more anterior so that together they form a broken x/ between the moderate-sized.11. its posterior border extensively free and deeply notched. distinct.126. . 1913. black-encircled. pupil vertical. Snout rather small. not particularly prominent. Body elongate.

148

NATIONAL MUSEUM BULLETIN

U.S.

distinctly narrower

axillary region

2

88

than greatest width of head;

adpressed, heel reaches beyond tip of snout; when
limbs are laid along the side, knee and elbow overlap appreciably;
when hind legs are bent at right angles to body, heels overlap some-

when hind

leg

is

No patagium. Skin of upper parts finely granular, a moderately
heavy glandular fold (parotoid) hangs behind the tympanum and angle
of jaw to about the level of the lower margin of jaw, a distinct (glandular?) flap extends out above the vent. Skin of throat and chest smooth,
that of belly and lower femur uniformly granular; no apparent traces
of a skinfold across the chest; no inguinal gland; no apparent vocal
sac. Skin of head not co-ossified with skull, roof of skull not exostosed.
Dimensions. Head and body, 62.7 mm.; head length, 19.2 mm.;
head width, 21.3 mm.; femur, 31.9 mm.; tibia (exclusive of dermal
flap), 33.7 mm.; foot, 42 mm.; hand 21.4 mm.
what.

A nearly uniform dorsal dark color, very sharply
Color in alcohol.
separated from the uniform (with exceptions noted below) ventral
light color; a series of black, narrow, finger-like projections extending
downward from the dark dorsal color into the ventral light color. The
dorsal dark color is now a deep blue and completely covers the top
of the head and back. There are margins of this dorsal color along
the margin of the lower jaw, along the upper surface of the forearm
and the lateral dorsal surface of the fourth finger; a narrow stripe of
this dorsal color extends from the hip along the dorsal margin of the
thigh to the knees. The dorsal surface of the shank bears the dorsal

ground

color, as

does the dorsal surface of the metatarsal region and the
is a pencil stripe along the distal

dorsal surface of the fifth toe. There
dorsal portion of the upper arm.
finger-like processes extend

A

series of

about half a dozen black,

down from the dorsal ground color
a similar number extend down both

be-

antween the head and groin;
teriorly and posteriorly from the stripe of dorsal color on the thigh,
on the inner (but not outer) margin of the shank, and several are
present on top of the foot, with a similar splash on the middle of each
fifth toe. The pale ventral ground color is interrupted only by some
brown markings on the ventral surface of the wrist and fourth finger
and by some brown spots slightly below and on each side of the vent,
on the ventral surface of the metatarsal region, the base of the fourth
toe, and the entire fifth toe.
The variation in the adult specimens is nearly imVariation.

perceptible. In the

two adults at hand,

BM 1902.5.27.26 [1947.2.24.22],

BM

the type, and
1913.11.12.126), the only perceptible difference
is that the type has a little more pronounced fold over the vent and a
little shorter snout. Color-wise, the two are exactly alike except that
the type lacks the pencil-like stripe along the distal, dorsal surface of
the upper arm.

FROGS OF COLOMBIA

— COCHRAN

AND COIN

149

Specimens Examined

COLOMBIA
Choc6:

BM

Condoto, Pena Lisa,
1913.11.12.126.
Rio Durango, northwestern Ecuador,

ECUADOR:
27],

BM

1902.5.27.26 [1947.2.24.

the type.

Agalychnis spurrelli Boulenger
Plate 23b
1913.

Agalychnis spurrelli Boulenger, p. 1024 (type locality, Pena Lisa, Condoto

1957.

Phyllomedusa

Choc6).— Dunn,

Diagnosis.

—An

1931, p. 415.

spurrelli.

— Funkhouser,

Agalychnis with

— Gorham,

1963, p. 24,

toe shorter

than the

1957, p. 39.

the

first

second, no dermal heel appendages, no pronounced glandular fold

above the vent although there is a glandular swelling, extensively
fingers, and a sloping snout. It differs from all Colombian
phyllomedusids except A. calcarifer in the presence of extensively
webbed fingers. It differs from A. calcarifer in lacking the two prominent triangular dermal heel appendages; in having distinct, clear
white, black-encircled, round spots scattered over the dorsum; in
having the palpebral membrane reticulate; in lacking the black,
finger-like projections from the dorsal ground color into the ventral
light surface; and in lacking any trace of pigment on the soles of the
hands and feet.
Description.—
1913.11.12.129 [1947.2.24.24], (cotype), from
Pena Lisa, Condoto, Choc6, Colombia. Vomerine teeth in two small,
slightly elongate series, the outer ends more anterior so that together
they form a very broad broken \/ between the moderate-sized, squarish choanae; tongue half as wide as mouth opening, elongated, its
posterior border extensively free and shallowly notched. Snout moderate, broadly triangular when viewed from above, somewhat sloping
in profile, the upper jaw not extending beyond lower; nostrils more
lateral than superior, very slightly projecting, half again as far
from eye as from tip of upper lip, separated from each other by an
interval equal to about two- thirds their distance from eye. Canthus
rostralis slightly defined; loreal region somewhat concave and somewhat oblique, the upper lip flaring out slightly below it. Eye moderate,
not particularly prominent, its diameter equal to its distance from

webbed

BM

nostrils; pupil vertical; interorbital distance half

again as great as

width of upper eyelid, equal to distance between nostril and eye,
upper eyelid not particularly broad, equal to distance between nostrils. Tympanum very distinct, about three-fifths the diameter of
eye, and practically in contact with it. Fingers extensively webbed,
web reaching nearly to base of disk on lateral side of the fourth
finger, fourth considerably longer than second, reaching to middle
337-262—70

-11

150

U.S.

of disk of third

NATIONAL MUSEUM BULLETIN

2

88

which covers nearly twice the tympanic area; no

projecting rudiment of a prepollex; an ulnar fold runs from elbow

along outer margin of forearm and hand to tip of fourth finger. Toes

webbed, the web on fourth toe reaching nearly to base of disk, web
other toes reaching to disk, third and fifth toes subequal, disk of
fourth toe covering half again the tympanic area; first toe shorter
than second, not opposable, a distinct oval inner but no apparent
fully

on

all

outer metatarsal tubercle; a well-developed tarsal ridge extending

from tip of fifth toe along outer margin of foot to heel; no dermal
appendage on heel. Body elongate, in postaxillary region a little
narrower than greatest width of head; when hind leg is adpressed,
heel reaches to tip of snout; when limbs are laid along the side, knee
and elbow overlap; when hind legs are bent at right angles to body,
heels overlap considerably. No patagium. Skin of upper parts very
finely glandular; a moderately heavy glandular fold (parotoid) hangs
behind the tympanum and angle of jaw to about the level of lower
margin of jaw; a pair of slightly projecting, rounded, glandular areas
just above the vent, but these not projecting out as a conspicuous
ridge or appendage; skin of throat and chest smooth, that of belly
and lower femur uniformly granular; no apparent traces of a skinfold across chest; no apparent vocal sac. Skin of head not co-ossified
with skull, roof of skull not exostosed.
Dimensions. Head and body, 92.8 mm.; head length, 28.2 mm.;
head width, 29.3 mm.; femur, 43.3 mm.; tibia, 48 mm.; foot, 63.2 mm.;

hand, 27.4

mm.

Color in alcohol.

now

—A rather somber colored frog for an Agalychnis,

a pale bluish above and pale yellowish below with no sharp line

demarcation between dorsal and ventral colors. The dorsal color
over the entire dorsal surface of head and body, in a narrow line
margining the lower lip, a line on the upper surface of forearm and
fourth finger, a trace of pigment on third finger, a narrow stripe along
the dorsal surface of thigh, a stripe along the margin of dorsal surface
of metatarsal region and on dorsal surface of fourth and fifth toes
(except on tip of fourth toe). The tarsal and ulnar folds are whitish,
and there are three very brilliant, clear white, black-encircled spots
on the back. The palpebral membrane is reticulated.
Variation.
The four specimens in the type series comprise two
adult females, an adult male, and a juvenile. The male differs in having
a well-developed, brown, horny pad on the underside of the thumb
and seems in general to be slightly more slender in build than are the
two adult females. The immature specimen (27.8 mm.) has faded to
a lavender rather than a pale blue color, but it is patterned like the

of
is

adults.

The

series of eight fresh

specimens

(CJG 2433-40) from

the Rio

.

FROGS OF COLOMBIA

Manso, Cordoba,

—COCHRAN

AND GOIN

151

by Fred Medem, adds

to our knowledge
breeding females in the series
have head and body lengths of 62.4 and 66.3 mm., while the males
range from 47.2 to 51.2 mm. in head and body length. Both females
and three of the males have the gaudy, rounded, black-bordered white
spots on the back as do those in the type series; and all the males
exhibit the brown patch on the inner side of the thumb, but this patch
is missing in both females.
Remarks. Boulenger (1913, pp. 1024, 1025) described and illustrated amplexus and the breeding process in this species. The leaf,
with the eggs attached, is still present with the type series.

collected

Two

of variation in this rare species.

Specimens Examined

COLOMBIA
Choc6:

BM

Condoto,

Pefia Lisa,

1913.11.12.127-30 (two specimens were

labeled as types in 1947 as 1947.2.24.24-5).

Cordoba:

Rfo Manso,

CJG

2433-40.

Genus Cerathyla Jiminez de
1871.

la

Espada

Cerathyla Jimfnez de la Espada, p. 63 (type species, Cerathyla bubalis

Espada)

A well-developed helmet but derm of head not
with skull; teeth or odontoids present on mandible and
palatines; expanded digital pads present; eye placed midway between
tip of snout and angle of jaw.
Generic diagnosis.

co-ossified

Key
a

1
.

to

Colombian Species of Cerathyla

Proboscis and superciliary processes in adults about 4

mm.

in

length;

tympanum

a2

separated from eye by its own greatest diameter (height)
rather than its least diameter (width)
C. proboscidea
Proboscis and superciliary processes in adults about 2 mm. in length;
tympanum separated from eye by its least diameter (width).
C. johnsoni

.

.

Cerathyla proboscidea Jiminez de la Espada

Plate 23f-h
1871.

Cerathyla proboscidea Jimfnez de la Espada, p. 64 (type locality not
given); 1875, pi. 5, fig. 2; 1899, p. 399.— Boulenger, 1882a, p. 453.—
Gadow, 1901, p. 211. Nieden, 1923, p. 360. Myers and Carvalho,

1945, p.
1945.

21.—Gorham,

Ceratohyla

cristata

Andersson,

Ecuador).—Gorham, 1963,
Diagnosis.

—A

1963, p. 21.
p.

18

(type

locality,

Rio

Pastaza,

p. 21.

Cerathyla with a long fleshy proboscis and elon-

gated, fleshy superciliary processes.

This species

may be

Colombian species

distinguished from C. johnsoni, the other

of the genus,

by having the proboscis and super-

152

U.S.

NATIONAL MUSEUM BULLETIN

2

88

about four millimeters long instead of about
2 millimeters long and by having the tympanum separated from the
eye by its height (long diameter) rather than by its width.
1910.7.11.71, from Rio San Juan, Choc6,
Description.
teeth in a pronounced V-shaped structure,
Vomerine
Colombia.
ciliary process in adults

—BM

anterior teeth in both series enlarged in a definite patch, palatine
teeth in two well-developed series nearly in contact medially and
passing laterally nearly to the maxillary; each mandible serrate along
entire length with fine, saw-toothed odontoids, with a much
enlarged odontoid at the tip of each ramus. No parasphenoid teeth.
Moderate-sized, rounded choanae lie in the triangle between the

its

vomerine and palatine teeth. Tongue about half as wide as mouth
opening, pear-shaped, its posterior border not free and not notched.
Snout arrow-head shaped, terminating in a 4-millimeter proboscis,
fleshy, rather flat and pointed in profile, the upper jaw extending

two millimeters

(exclusive of proboscis)

beyond lower;

nostrils

more

superior than lateral, not projecting, their distance from end of snout
about equal to that from eye, separated from each other by an interval

equal to about one-half their distance from eye. Canthus rostralis
sharply defined but low; loreal region very slightly concave and
very oblique, the upper lip flaring out strongly below it. Eye small,
not prominent, its diameter equal to its distance from nostril; interorbital distance more than twice the width of upper eyelid (exclusive
of superciliary appendage), which is slightly greater than distance

between

nostrils.

Upper

eyelid with a fleshy superciliary

appendage

four millimeters long. Palpebral membrane not reticulate. Tympanum
very distinct, twice as high as wide, its long diameter about equal to

diameter of eye, separated from eye by a distance nearly equal to
long diameter. Fingers with a trace of web at base, fourth finger
considerably longer than second, reaching to disk of third, which is
not so wide as the narrow width of the tympanum; no projecting
rudiment of a pollex; ulnar ridge well developed. Toes about one-fourth
webbed, the web on fourth toe reaching the middle of the preantepenultimate phalanx, third and fifth toes subequal, disk of fourth
its

toe covering an area about one-half the diameter of disk of third
finger; a distinct oval inner

and a smaller but equally

distinct

rounded

outer metatarsal tubercle; a well-developed tarsal ridge; a dermal
appendage on heel. Body not elongate, in postaxillary region much

narrower than greatest width of head; when hind leg

is

adpressed,

when limbs are laid along the side, knee and
considerably; when hind legs are bent at right angles

heel reaches tip of snout;

elbow overlap
to body, heels overlap considerably. No patagium. Skin of upper
parts very finely granular, but the projecting neural spines of the
dorsal vertebrae give the appearance of a median row of well-


FROGS OF COLOMBIA

— COCHRAN

AND GOIN

153

developed warts on the back. The lateral margin of postorbital process
of helmet makes a shelf above the tympanum; skin of throat and
chest thin and smooth, that of belly and lower femur very thin and
very finely granular; no skinfold across chest; no inguinal gland;
no apparent vocal sac. Skin of head not co-ossified with skull, roof
of skull exostosed and forming a well-developed casque. The helmet
is triangular in shape along the margin of jaws, the can thai ridges
curve and meet between the nostrils; above, the helmet terminates
posteriorly in two well-developed postorbital spines with a curving
posterior margin between them. On the side of the head the helmet
behind the eye and between the angle of jaw and the postorbital
spine is interrupted only by the tympanum.
Dimensions.

—Head and body, 44.4 mm.; head length

mm.; head width, 25.5 mm.; femur,
25 mm.; heel-to-toe, 36.1 mm.; hand, 16 mm.
proboscis), 20.7

(exclusive of

22.5

mm.;

tibia,

—A very pale tan frog with a

little dark brown along
above and behind the eyes, and above the
sacrum and above the tip of the urostyle. There are faint traces of
several narrow, darker stripes above the legs.
Remarks. The described specimen was the only one examined.

Color in alcohol.

the canthal ridge, a

little

Cerathyla johnsoni Noble

Plate 23c-e
1917.

Cerathyla johnsoni Noble, p. 798, pi. 37 (type locality, Colombia, Antioquia, 14 mi. north of Mesopotamia, Santa Rita Creek) 1926a, p. 18.
;

Dunn, 1944c,
1945.

p. 517.

Cerathyla bubalis.

— Myers and Carvalho,

1945, p. 21.

— Gorham,

1963, p.

21.

—A

mm.) proboscis and
and with the tympanum separated from the
eye by a distance about equal the width of the tympanum.
Diagnosis.

Cerathyla with short (about 2

superciliary processes

From C. proboscidea, the only other member of the genus now
known from Colombia, it can be distinguished by the shorter proboscis
and superciliary processes (2 mm. instead of 4 mm.) and by having
its tympanum separated from the eye by a distance equal to its
width rather than

its

height.

— CNHM

63850, from eight kilometers south of
Colombia. Vomerine teeth in a pronounced
V-shaped structure, anterior teeth in both rami of the V are enlarged
Description.

Valdivia,

Antioquia,

into a definite patch, palatine teeth in

two well-developed

series

nearly in contact medially and passing laterally nearly to the maxillary;

each mandible serrate along its entire length with fine, saw-tooth
odontoids with a much enlarged odontoid at the tip of each ramus.

154

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parasphenoid teeth. Moderate-sized, somewhat ovate choanae

lying in the triangle between the vomerine and palatine teeth. Tongue
about one-third as wide as mouth opening, rounded, its posterior

border fused and unnotched. Snout arrow-head shaped, terminating
about two millimeters in length, rather

in a small fleshy proboscis

and pointed in profile, the upper jaw extending about two millimeters (exclusive of proboscis) beyond lower; a short, fleshy appendage
at the base of each jaw; nostrils more superior than lateral, somewhat
projecting, their distance from end of snout about equal that from
eye, separated from each other by an interval equal to about one-half
their distance from eye. Can thus rostralis sharply defined but low;
loreal region very slightly concave and very oblique, the upper lip
flaring out strongly below it. Eye small, not prominent, its diameter
equal to its distance from nostril; interorbital distance equal to about
twice the width of upper eyelid, which is slightly greater than distance
flat

between nostrils. Upper eyelid with a fleshy superciliary appendage
about one millimeter long. Palpebral membrane not reticulate.
Tympanum very distinct, nearly twice as high as wide, its long
diameter one-third again as great as diameter of eye, separated from
eye by a distance nearly equal to its width. Fingers with but a trace
of web at base, fourth finger considerably longer than second, reaching
to disk of third, which is not so wide as the narrow width of the
tympanum; no projecting rudiment of a pollex; ulnar ridge well
developed. Toes about one-fourth webbed, the web on fourth toe
reaching the middle of the preantepenultimate phalanx, third and
fifth toes subequal, disk of fourth toe covering an area about one-half
the diameter of disk of third finger; a distinct oval inner and a smaller
but equally distinct rounded outer metatarsal tubercle; a welldeveloped tarsal ridge; a well-developed dermal appendage on heel
and another on knee. Body not elongate, in postaxillary region much
narrower than greatest width of head; when hind leg is adpressed,
heel reaches tip of snout; when limbs are laid along the side, knee
reaches to axilla; when legs are bent at right angles to body, heels
overlap considerably. No patagium present. Skin of upper parts
very finely granular, but the projecting neural spines of dorsal vertebrae give the appearance of a median row of well-developed warts on
the back. The lateral margin of postorbital process of helmet makes a
shelf above the tympanum. Skin of throat and chest thin and smooth,
that of belly and lower femur very thin and very finely granular; no
skinfold across chest; no inguinal gland; no vocal sac apparent.
Skin of head not co-ossified with skull, roof of skull exostosed and
forming a well-developed casque. Helmet triangular in shape along
margin of the jaws, can thai ridges curve and meet between the
nostrils; above, the helmet terminates posteriorly in two well-

.

FROGS OF COLOMBIA

—COCHRAN

AND GOLN

155

developed postorbital spines with a curving posterior margin between
them. On the side of the head, the helmet behind the eye and between
the angle of the jaw and the postorbital spine is interrupted only by
the

tympanum.

Dimensions.

—Head

and body 45.5 mm.; head length 22.4 mm.
head width, 23.3 mm.; femur, 23.6 mm.;

(exclusive of proboscis);

mm.; hand, 15.6 mm.
very pale tan frog with a darker brown triangular area on the back just posterior to helmet, the skin above the
tips of neural spines dark brown, a small brown spot above the middle
of urostyle, a dark triangular area below vent, and chin, throat, and
chest dark chocolate brown. The brown on the chest extends onto the
undersurface of upper arm and then crosses as a band across the upper
surface of forearm. The margins of both upper and lower lips are
mottled with dark brown. Three narrow, rather pale, brown crossbands occur on upper surface of each thigh, shank, and foot. Dorsal
tibia, 26.5

mm.;

heel-to-toe, 34.7

Color in alcohol.

—A

surfaces of fingers are rich chocolate

brown

in contrast to the pale

dorsal surface of the hand.

Remarks. We cannot follow Myers and Carvalho in synonymizing
form with C. bubalis. We strongly suspect that time and material
will demonstrate rather extensive geographic variation in this group
and that many of the nominal forms will prove to be geographic
races of one or two widespread polytypic species.
this

Specimens Examined

COLOMBIA

AMNH 1341
CNHM 63850.

Antioquia: Santa Rita Creek, 14 mi. north of Mesopotamia,
(type), 1342-3 (photographs); eight km. south of Valdivia,

Genus Cryptobatrachus Ruthven
1916.

Cryptobatrachus Ruthven,
boulengeri

1916b,

p.

1,

(type species,

Cryptobatrachus

Ruthven)

Vomerine teeth in two nearly straight transverse
which nearly touch on the midline and which lie behind the
rounded choanae; sacral diapophyses nearly rounded; female carries
the eggs on her back; eggs hatch directly into frogs; a single subgular
vocal pouch in the male.
Generic diagnosis.

series,

Key
a

Without

1
.

to Colombian Species of Cryptobatrachus

distinct pale light spots

on dorsum and with thighs dark with

light

blotches.
b1 .

Distance from anterior corner of eyelid to naris distinctly less than distance
from anterior to posterior corner of eyelid; dorsum rough and dorsolateral
folds fairly distinct

C. fuhrmanni

156
b2

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Distance from anterior corner of eyelid to naris about equal to that from
anterior to posterior corners of eyelid;

dorsum

fairly

smooth and dor-

solateral folds not so distinct

a2

.

With

distinct pale light spots

C.

boulengeri

on the dorsum and both pale thighs marked

with three or four distinct dark crossbands

C.

nicefori

Cryptohatrachus nicefori, new species

Figure


Diagnosis. —A

7

MLS 138, an adult(?) female collected at La Salina,
Holotype.
Boyaca, Colombia, by Hno. Niceforo Maria.
spicuously

Cryptohatrachus with

marked with

thin,

Hyla-\ike

thighs

con-

three or four distinct, broad, black cross-

bands, and with distinct pale (yellow in preservative) scattered light
spots on the dorsum.

may be distinguished at a glance from both
fuhrmanni by the thin thighs and by the striking
pattern on thighs and dorsum.
Description of holotype. Vomerine teeth in two long, transverse
series, lying close together behind the posterior borders of the moderate-sized, somewhat ovate choanae; tongue three-fourths as wide
as mouth opening, broadly rounded, its posterior border fused and
This

new

C. boulengeri

species

and

C.

Figure

7.

Cryptohatrachus nicefori,

mouth

parts,

holotype,

which are

MLS
X 2).

138

(same

size,

except

FROGS OF COLOMBIA

— COCHRAN

AND GOIN

157

not notched. Snout moderate, somewhat triangular when viewed
from above, slightly rounded in profile, the upper jaw extending
slightly beyond lower; nostrils more lateral than superior, very
slightly projecting, their distance from end of snout about one-half
that from eye, separated from each other by an interval equal to
about their distance from eye; distance from nostril to anterior
corner of eye about equal to distance from anterior to posterior
corners of eye.

Can thus

rostralis well defined; loreal region slightly

concave and nearly vertical, the upper lip flaring out but very slightly
below it. Eye moderate, protruding, prominent, its diameter equal
to its distance from nostril; palpebral membrane not reticulate;
interorbital distance as great as width of upper eyelid, which is
relatively wide and slightly greater than distance between nostrils.
Tympanum distinct, about one-third the diameter of eye, separated
from eye by a distance nearly equal to its own diameter. Fingers with
a trace of web between second and third and between third and
fourth, fourth finger about one-half disk longer than second, reaching
to disk of third, which just covers the tympanic area; no projecting
rudiment of a pollex; no ulnar ridge. Toes slightly more than one-half
webbed, the web on fourth toe reaching to base of the antepenultimate
phalanx, third toe slightly longer than fifth, disk of fourth covering
about two-thirds the tympanic area; a distinct oval inner but no
apparent outer metatarsal tubercle; no tarsal ridge; a wart-like dermal
appendage on heel. Body moderately elongate, in postaxillary region
distinctly narrower than greatest width of head; when hind leg is
adpressed, heel reaches eye; when limbs are laid along the side, knee
and elbow just meet; when hind legs are bent at right angles to body,
heels overlap greatly; thighs thin and Hyla-\ike. No patagium. Skin
of upper parts smooth; no structural dorsolateral folds evident; a
rather narrow glandular ridge passing above tympanum; skin of throat
and chest smooth, that of belly and lower surface of thigh uniformly
granular; no apparent traces of a skinfold across chest; adult (?)
female, no vocal sac apparent. Skin of head not co-ossified with skull,
roof of skull not exostosed.
Dimensions. Head and body, 39.3 mm.; head length, 14.6 mm.;
head width, 14 mm.; femur, 17.3 mm.; tibia, 21.1 mm.; heel-to-toe,
29.4 mm.; hand, 13.4 mm.
Color in alcohol.
Dorsum dark gray with scattered, distinct,
more or less rounded yellow spots. These spots range from one-half
to two-thirds the diameter of tympanum. Between axilla and groin
are a series of rather distinct in some cases nearly vertical
black
blotches. Each thigh is crossed by three distinct, broad, black bands
which extend down onto the anterior and posterior faces of the thigh
but which fail to meet on the undersurface of the thigh. Paler but

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NATIONAL MUSEUM BULLETIN 288

somewhat similar crossbands are on the dorsal surfaces of the shanks
and arms. The specimen is rather dark below.
Remarks. At first glance, the thigh pattern of this species makes
one think of Hyla boulengeri; in fact, this species has a more Hyla-\ike
mien than has either C. boulengeri or C. fuhrmanni.
The vomerine teeth and the more or less cylindrical sacral di-

new

species should be described as a
completely unknown.
The holotype, which is in the collection of the Museo del Instituto
de La Salle, may well be the same specimen on which Dunn (1943,
p. 311) based his statement that there was an undescribed species of

apophyses convinced us that
Cryptobatrachus.

The

life

this

history

is

Cryptobatrachus in northern Colombia.

The holotype was

the only specimen examined.

Cryptobatrachus fuhrmanni (Peracca)

Figure 8
1914.

Hyla fuhrmanni Peracca,

p.

108 (type locality, Colombia, Guaca [=Heli-

conia, Antioquia], 1,600 m.).

1917.
1926.

33.— Goin,
1944.

Dunn, 1943, p. 310.
Noble, 1917, p. 805.
Hyloscirtus fuhrmanni.
Cryptobatrachus incertus Barbour, p. 193 (type locality, Colombia, Antioquia, Sons6n).
Barbour and Loveridge, 1929, p. 245. Liu, 1935, p.
1961, p.

8.— Gorham,

Cryptobatrachus fuhrmanni.

1963, p. 21.

— Dunn, 1944c,

p.

517 [1957,

p. 91].

Diagnosis. A Cryptobatrachus with thick thighs that are rather
dark posteriorly.
This species may be distinguished from C. nicefori by the lack of
distinct dark crossbands on the thighs and distinct light spots on the
back. From C. boulengeri it may be distinguished by its having a
rougher, wartier skin, generally darker color both above and below,
more distinct folds along the dorsolateral region, and the distance
from the anterior corner of the eyelid to the naris distinctly less than

the distance between the anterior and posterior corners of the eyelids.
Description.
11616 (type of incertus) from Sons6n, Antioquia,
Colombia. Vomerine teeth in two short, heavy, transverse series lying

— MCZ

close together (but with a distinct gap) on a level with the posterior
borders of the choanae; tongue narrow, about half as wide as mouth
opening, its posterior border fused and without a distinct notch. Snout
moderate, broadly ovate when viewed from above, slightly projecting
when viewed in profile, the upper jaw extending slightly beyond lower;

no s trils more lateral than superior, very slightly projecting, their distance from end of snout about one-half that from eye, separated from
each other by an interval about equal to their distance from eye.
Can thus rostralis well denned; loreal region concave and somewhat
oblique; the upper lip flaring out slightly below it. Eye moderate,

FROGS OF COLOMBIA

— COCHRAN

AND GOIN

159

diameter equal to its distance from nostril interabout one-half the width of upper eyelid, which is
about equal to distance between nostrils. Tympanum very distinct,
about one-half the diameter of eye, separated from eye by about

very prominent,

its

;

orbital distance

one-half

its

own

diameter. Fingers entirely free, fourth finger con-

siderably longer than second, reaching to middle of disk of third which

more than covers the tympanic area; no rudiment of a pollex.
more than one-half webbed, web to base of disk of
second and fourth toes, third and fifth toes subequal, disk of fourth

slightly

Toes

slightly

toe covering the tympanic area; a distinct oval inner and a smaller,

rounded outer metatarsal tubercle; no tarsal ridge; no
dermal appendage on heel. Body not elongate, in postaxillary region
but little narrower than greatest width of head; when hind leg is
less-distinct,

adpressed, heel reaches beyond naris;

when limbs

are laid along the

knee and elbow considerably overlap, the knee reaching the
axilla; when hind legs are bent at right angles to body, heels slightly
overlap. No patagium. Skin of upper parts glandular, with prominent,
elongate, wart-like tubercles arising from various areas in which the
glandules of the skin appear particularly concentrated, these tubercles
particularly conspicuous around the tympanum and at the bases of
side,

Figure

8.

Cryptobatrachus fuhrmanni,

CNHM 63892

(same

size).

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from behind the eye, above the
Another row beginning above the
axilla and extending posteriorly to between the sacrum and coccyx,
thus forming a sort of dorsolateral fold; a few less well-developed
tubercles on upper surfaces of arm; skin of throat, chest, belly, and
lower femur uniformly and very finely granular; no skinfold across
the jaw.

tympanum

distinct

row

of tubercles

above the

to

axilla.

chest; vocal sac not apparent.

Dimensions. Head and body, 46.6 mm.; head length, 17 mm.;
head width, 17.3 mm.; femur, 27.7 mm.; tibia, 28.9 mm.; foot, 36.7

mm.; hand,

16.4

mm.

A very dark frog. Entire dorsum, top of head,
and upper surfaces of limbs a chocolate brown; under magnification
this brown can be seen to be mottled faintly with an even darker
brown. Undersurfaces of chin, throat, and belly a pale brown with
the undersurfaces of the thighs a mottling of the dorsal and ventral
ground colors. Upper lip dark with four vertical bars of a lighter hue;
two of the bars enter the eye.
There are too few specimens available to make an
Variation.
adequate study of variation, but from those that are available it
appears that C. juhrmanni is a rather uniform species. All the male
specimens are rather dark and rather rough, with fairly well-developed folds along the dorsolateral region.
A large female (CNHM 82025, with the data "Colombia") seems
to represent this form. This frog, measuring 60.7 millimeters in head
and body length, was carrying eggs when captured. One of the eggs
still with this female contains an embryo with pigment and welldeveloped eyes. There are 24 egg pockets in the skin on her back.
The pattern on the rear of her thighs is paler than in the examined
males of this form, but her short snout and the roughness on the top
of her head seem to indicate that this female should be identified as
juhrmanni rather than boulengeri or nicejori.
Remarks. The name "juhrmanni" has had a somewhat checkered
Color in alcohol.

history. Peracca (1914, p. 108) described the frog in the genus Hyla.

Noble (1917, pp. 804, 805), surely without ever having seen a HyloCryptobatrachus boulengeri (which had been described
by Ruthven in 1916) to the genus Hyloscirtus, along with Hyla juhrscirtus, assigned

manni

of Peracca. Since

we have now

adults of both sexes and the
species of Cryptobatrachus,

seen recent material of the

young

of Hyloscirtus, as well as all three

we can

categorically state that the two

genera are distinct and that Hyloscirtus has nothing to do with the
problem under discussion. Ruthven (1922, p. 551) took issue with

Noble about assigning Cryptobatrachus
his

own

species, boulengeri, to the

but he assigned
juhrmanni. Barbour

to Hyloscirtus

synonymy

of

Three distinct populations of Cryptobatrachus are now known from Colombia. p. from boulengeri San Gil in Santander are the only ones of that species recorded from outside the Santa Marta Mountains. Peracca's name antedates Barbour's by twelve years. 193). 1917. 1922. and the data.FROGS OF COLOMBIA — COCHRAN AND GOIN 161 (1926. 1961. 538. and for it two names are available: fuhrmanni Peracca with the type locality "Guaca" (now known as Heliconia). San Lorenzo. p. —Hellmich. — p. 1.— Gorham. 21. 311) indicated that there was an undescribed Cryptobatrachus in northern Colombia. The third population is in the northern end of the Cordillera Oriental of the Andes of northern Colombia in the states of Santander and Boyacd. 1916b. 1952. described Cryptobatrachus incertus from Sonson in Antioquia. One is from the Sierra Nevada de Santa Marta in Magdalena. 1944c. Hyloscirtus boulengeri. 1939.— Dunn. Colombia). apparently for the first time. 1916b. 517) (1943. about 70 kilometers south-southeast of Medellin. Bellavista. They certainly are not identical with nicefori of adjacent Boyacd. Magdalena. Peters. MCZ 11616 (type). p. The specimens Specimens Examined COLOMBIA Antioquia: Jeric6. and he (Dunn. 517. — Ruthven. p. 1917. 16266. Another is in the Cordillera Central of the Andes of northern Colombia in the state of Antioquia. Barbour and Loveridge. 12 Hy la fuhrmanni. for which the name boulengeri Ruthven (type locality. Sonson. 245. pointed out correctly. p. — p. 1916. with the type locality Sons6n. 55. (type locality. 1944c. 2. and incertus Barbour. p. apparently without checking type localities. Goin. p. Cryptobatrachus fuhrmanni. 805. 8. MCZ 24887. p. p. p.— J. above Rio Porce. No precise locality: 82025. Cryptobatrachus boulengeri Ruthven. 63892. 1963. . San Lorenzo) is available. (MZUM 74812 and MLS 202) listed under C. CNHM CNHM Cryptobatrachus boulengeri Ruthven Plate 24a-c 1916. 1 pi. 1922. p. 1929. Ruthven. if correct. would seem to indicate that boulengeri probably occurs that throughout the little-collected Sierra de Perijd and Sierra de Ocafia form the northern extension of the Oriental Andes. that the name Cryptobatrachus fuhrmanni applied to the population in the Cordillera Central and the name Cryptobatrachus boulengeri applied to the one in the Sierra Nevada de Santa Marta. — — Noble. for that population we herein propose the name Dunn nicefori. about 18 kilometers southwest of Medellin.

Tympanum very distinct. nicefori by the lack of very distinct crossbands on the thighs and light spots on the back. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. separated from eye by a distance equal to about one-half its own diameter. traces of a skinfold across the chest. slightly projecting. transverse series lying close together on a level with the posterior borders of the small.162 U. NATIONAL MUSEUM BULLETIN 288 Diagnosis. roof of skull not exostosed. fourth finger considerably longer than second. Vomerine teeth in two short. no tarsal ridge. Toes slightly more than one-half webbed. Snout moderate. no inguinal gland. From C. no dermal appendage on heel. both above and below. skin of throat and chest smooth. the snout extending slightly beyond lower jaw. broadly triangular when viewed from above. nostrils more lateral than superior. This species may be distinguished from C. in postaxillary region a little phalanx. third and narrower than greatest width of head. No patagium. not one-fourth greater than its about equal to width of upper eyelid. and by having the distance from the anterior corner of the eyelid to the naris about equal the distance between the and posterior corners of the eyelid. when limbs are laid along the side. heels just touch. loreal region slightly concave and somewhat oblique. tongue half as wide as mouth opening. by being generally paler. the web on fourth toe reaching the base of the penultimate fifth toes subequal. from San Lorenzo. no projecting rudiment of a pollex. the upper lip flaring out moderately anterior Description. Can thus rostralis fairly well defined. heel reaches beyond tip of snout. Colombia. knee reaches into axilla. rounded in profile. . vocal sac not apparent. their distance from end of snout about one-half their distance from eye. below it. Magdalena. rounded choanae. disk of fourth toe just covering the tympanic area. when hind legs are bent at right angles to body. which is relatively wide and about equal to the distance between nostrils. by having much less distinct dorsolateral folds. which snugly covers the tympanic area. when hind leg is adpressed. about one-fourth the diameter of eye. broadly cordiform. just reaching to middle of disk of third.—A Cryprobatrachus with thick thighs that are faintlymottled posteriorly. Fingers without webs at base. a rather narrow glandular ridge encircling upper part of tympanum. fuhrmanni it may be distinguished by its smoother skin. 48530 (holotype).S. interorbital distance Eye moderate. its diameter distance from nostril. Skin of head not co-ossified with skull. separated from each other by an interval equal to about their distance from eye. that of belly and lower femur uniformly and very finely granular. Skin of upper parts smooth. —MZUM particularly prominent. Body not elongate. its posterior border very slightly free and very shallowly notched.

54623-6. The crossbands on the hind legs seem to be more distinct than they are in C. on road to San Lorenzo.FROGS OP COLOMBIA — COCHRAN AND GOIN 163 — Dimensions. MLS 202. 12. the ventral margin. Santa Marta Mts. However. broader body.. No structural differences can be observed in the Variation. 10 mm. series of fresh . Quebrada Viernes Santo. head length. 19. In some of the specimens.7 mm. 48531-5. Santandeb: San Gil. Santa Marta Mts.1.1 mm. Mts. 33. San Lorenzo. 48530 (holotype). A similar sort of light-bordered. and another dark band extends from the eye backward through the tympanum. Specimens Examined COLOMBIA Magdalena: Santa Marta MZUM MZUM BM Mts. 25. these specimens are dark gray on the chin becoming lighter posteriorly. nicefori) but this may be due to the fact that the specimens of boulengeri are fresher than those of fuhrmanni.8 mm. Santa Marta Mts.. p. specimens from the Rio Minca in the Sierra Nevada de Santa Marta. to a lesser extent. 118734. Color in alcohol.1. Santa Marta 8980. head width. foot. Cincinnati. while the undersurfaces of the thighs and. —Medium.7 mm. palatines. one of the most conspicuous pattern elements is a broad dark band between the eyes bordered with a very pale narrow margin.. Quebrada Hueva Santa. Structurally. Head and body. this band has a straight margin anteriorly and a v-shaped margin posteriorly so that the interocular bar has the form of an inverted triangle. 12.25. 47408-9. 48614-5. In each of the fresh specimens. 48617.3 mm. MCZ USNM MZUM MZUM MZUM MZUM MZUM Genus Gastrotheca Fitzinger 1843. tibia. Generic diagnosis. Ventrally.. fuhrmanni (though not so marked as they are in C. Santa Marta Mts. permits a little better understanding of the pattern variation in this species than we have had heretofore. Quebrada. 54627-8. but not the dorsal margin. Rfo Minca. hand.to Hyla marsupiata Dum€ril and large-size hylids without teeth on the mandible. or parasphenoid.. collected by Fred Medem. above and a — This specimen dirty. 30 (type species. 19.. Gastrotheca Fitzinger. 54638.. amount of webbing on the feet. 74812. of the latter band is bordered by white. the specimens of boulengeri seem to have a relatively shorter. 54629. dark band extends downward and slightly anteriorly from each eye to the margin of the upper lip. 54622. 54631-2.3 mm... 48300. 48537-8. femur. San Lorenzo. the posterior portion of the belly and the undersurfaces of the shanks and arms are bright yellow. yellowish —A now is a warm medium brown brown below without a trace of pattern.. 54630. 2452-3. CJG 2341-50. with a well-defined pouch . 1921.. whereas the specimens of fuhrmanni seem to have heavier thighs. Bibron). 54620-1.

aureomaculatus d2 . Diagnosis. NiceToro Maria in 1963. dermal appendages on heels G. collected by Hno. . with dermal appendages on the heels. Dorsal pattern bright and conspicuous. nicefori tip of urostyle c2 Tibia length not more than one-half the distance from snout to tip of G. intersquamosal shelf absent or incomplete but not forming a nearly straight margin at back of skull. Gastrotheca bufona. vomerine teeth in two short series. an adult male from Ventanas (about 50 kilometers [by road] northwest of Yarumal) Antioquia. medemi urostyle Cranial derm free from skull. Cranial derm co-ossified with roof of skull. monlicola argenteovirens what mottled c2. intersquamosal shelf complete and forming a nearly straight posterior margin to skull. derm of the head may be co-ossified with the skull or it may be free. d 1 . . adult male with a rudimentary pouch on the back and a single. and with the derm of the head co-ossified with the skull.. small. Key a to Colombian Species of Gastrotheca Superciliary processes forming distinct triangular appendages on the upper 1 . Cranial derm co-ossified with roof of skull. a2 . bufona Cranial derm not fused with roof of skull. opening posteriorly either by a round.) and belly generally someG. helenae Dorsal pattern somber or plain. b2. new species Figure 9 — Holotype. but the pouch never a slit-like trough as in Nototheca. cornutum tiny or absent Without superciliary processes forming distinct triangular appendages on upper eyelids. females 70+ mm. b1 . in which case the roof of the skull is exostosed. dermal apG. Tibia length distinctly more than one-half the distance from snout to c G. Colombia.S. A Gastrotheca with the upper eyelids bearing triangular fleshy appendages. Dorsum dark with numerous. eyelids. c1 . — . unpaired vocal pouch. 6 1 . This new species can be distinguished at a glance from all Colombian Gastrotheca except cornutum by the well-developed superciliary dermal process. well-developed pendages on heels b2. Dorsum pale with a very conspicuous dark mid-dorsal stripe G. rounded light spots G. NATIONAL MUSEUM BULLETIN 2 88 on the back of the female. 1 .) and belly not G. d1 Size large (males 60-70 mm. .164 U. MLS 344. It differs from cornutum in having the derm of the head coossified with the skull. puckered aperture or by a longitudinal slit. mertensi generally mottled d2 Size smaller (generally less than 60 mm.

— FROGS OF COLOMBIA Figure 9. 12 AND GOIN MLS 344 (same 165 size). holotype. 337-262—70 — COCHRAN Gastrotheca bufona. .

Without a flat. no projecting rudiment of a pollex. lying close together between the posterior borders of the small. elongate dermal appendage on heel. Fingers un webbed. heel reaches slightly beyond tip of snout. the upper jaw extending hardly at all beyond lower. fleshy ridge passes from behind the eye above the to the point of origin of the soft dorsolateral fold. rounded. no trace of a skinfold across chest. heels overlap greatly. separated from eye by a distance nearly equal to its own long diameter. no vocal sac apparent plate forming a bony intersquamosal margin at the back of the skull. no distinct tarsal ridge. toe. when hind legs are bent at right angles to body. reaching to middle of disk of third. Skin of throat. when limbs are laid along the side. Skin of upper parts coarse and glandular. — Head and body. NATIONAL MUSEUM BULLETIN Description oj holotype.. head width.3 mm. 166 U. triangular. Tympanum distinct but inconspicuous. warty. their distance from tip of upper jaw about equal that from eye. the dorsolateral folds well developed anteriorly projections. belly. warty. in postaxillary region distinctly narrower than greatest width of head when hind leg is adpressed.. its long diameter about one-half the diameter of eye.5 mm. a low but distinct. a well-developed. tongue two-fifths as wide as mouth opening. nostrils lateral.: heel-toDimensions. Skin of co-ossified with skull.. 23. "horns". and unnotched. loreal region concave and nearly vertical. not projecting. —Vomerine teeth in 2 88 two rounded patches. Upper No patagium. eyelids bearing triangular. head length. Eye moderate. . Upper distance slightly eyelid developed into an elongate. no distinct outer tubercle. rounded when viewed from above. not prominent. Snout moderate.3 mm. a distinct but minute oval inner metatarsal tubercle. rounded choanae. Body somewhat elongate for a Oastrotheca. its posterior border free membrane not palpebral reticulate. tympanum A bony and bearing white.S. 25.2 mm. fourth finger considerably longer than second..8 mm. disk of fourth half again as large as the tympanic area.. 37. warty ulnar ridge. tibia. hand. 55.0 mm. and lower surface of thigh uniformly granular.. 20. externally.6 mm. roof of skull exostosed. truncate in profile. Toes webbed at base between third and fourth and fourth and fifth. Third toe distinctly shorter than fifth. pointed superciliary process. chest. its diameter one-third again as great as its distance from nostril. 51. transverse. nearly straight head Adult male. which is twice the size of the tympanic area. Can thus rostralis sharply defined. knee and elbow overlap considerably. femur. greater than width of upper eyelid at its interorbital base. web not quite reaching base of antepenultimate phalanx of fourth toe. separated from each other by an interval equal to about two-thirds their distance from eye. 31.

— Superficially the holotype looks more like G. 1963. Ecuador. from Rio San Joaquin. This species differs from all Colombian Gastrotheca except bujona having well-developed triangular processes growing out of the upper eyelids. Description. Distinct bands of dark pigment on a lighter ground color cross the dorsal surfaces of the arms and hands and the legs and feet. tongue two-thirds as wide as mouth opening. these dark bands being rather conspicuous on a ground color of light brown. 1963. black spots in a row along the margin of the upper jaw. Larger.FROGS OF COLOMBIA Color in alcohol. 54718. — Gorham. triangular dermal appendages. and in having well-developed dorsolateral folds that bear tiny. p. — COCHRAN AND GOIN 167 —A gaudy frog with a bright pattern on a ground and top of head and loreal region with rounded black dots. 1898a. Remarks. rounded. in reference to the frog's gaudy appearance. Cauca. Sides scattered. Pacific side. Gastrotheca cornutum (Boulenger) Figure 10 1898. The margin of the lower lip is white but the chin and underside of the throat are coal black. Vomerine teeth in two small. 124 (type locality. broadly ovate. This remarkably distinct species is known from the single specimen collected by Hno. distinct." a jester or clown.— Nieden. p. p. 321. 188. ceratophrys of Panama than any other form. p. 1901. Dorsolateral folds nearly black but the tiny. but it differs from that species in having dermal appendages on the heels. Cachabe). in having the skin of the cranium co-ossified with the skull. Niceforo Maria at Ventanas. 21. 1904. A Gastrotheca with superciliary processes but with dermal appendage on heel reduced or absent. triangular fleshy processes they bear are nearly white. It differs from bujona in the reduced dermal appendages on the heels and in having the cranial derm free from the skull.500 meters. 1. Colombia. — Diagnosis. p. The specific name is from the Spanish "bufon. oval series lying close together between the small. rounded choanae. color of tannish brown. Transverse bands of dark brown cross the dorsum from the back of the head to hind end of the body. its in — CNHM . Nototrema cornutum Boulenger. Gastrotheca cornutum. 40. The chest and belly and the underside of the arms and legs are yellowish and heavily mottled with dark brown and black areas. margin of upper jaw between these spots nearly white. The cranial derm is not fused to the skull. 1923.—Peracca. This is another of the many new species of frogs from Colombia for which our science is indebted to our good friend Hermano Niceforo Maria.— Gadow.

web reaching middle of antepenultimate phalanx of fourth toe. definitely greater than distance between nostrils. but no distinct outer. fourth finger considerably longer than second. NATIONAL MUSEUM BULLETIN 2 88 and not notched. which more than covers the tympanic area. Fingers with a trace of web at base. interorbital distance about equal the width of upper eyelid. its diameter equal to its distance from nostril. Body moderately elongate for a Gastrotheca. CNHM 63892 (same size). the upper jaw extending slightly beyond lower.third that from eye. Can thus rostralis well defined.— 168 U. third and fifth toes sub- tympanic area. a distinct oval inner. nostrils more lateral than superior. Snout short. which is relatively wide.S. no dermal appendage on heel. not projecting. no tarsal ridge. loreal region concave and very oblique. ovate when in profile. reaching to disk of third. posterior border not free viewed from above. about one-third the diameter of eye. equal. in postaxillary region definitely narrower than greatest width of head. Toes slightly more than one-half webbed. their distance from end of snout about one. separated from each other by an interval equal to about two-thirds their distance from eye. disk of fourth toe covering the Figure 10. sloping Tympanum moderately distinct. Gastrotheca cornutum. prominent. Eye moderate. the upper lip flaring out below it. an external swelling indicating a rudiment of a pollex. . metatarsal tubercle. separated from eye by a distance nearly equal to twice its own diameter.

in having knee reach to axilla (to elbow in the type) in having seven definite rows of warts transversely across back (absent in the type) and in being in general a more slender frog.. a rather narrow glandular ridge encircles upper part of tympanum. when limbs are laid along the side. 27. cornutum. and along the median line of the back to the upper margin of the egg sac opening.FROGS OF COLOMBIA — COCHRAN AND GOIN 169 when hind leg is adpressed. wart-like structures in inconspicuous transverse rows across the back.4 mm. Entire ventral surface dark but not so dark as the dorsum. in head-body length as against 76. below this lateral stripe. alcohol. skin heavily folded along sides.. The type is a pale frog with tannish yellow dorsal ground color. 30.. posteriorly between the eyes and parietal ridges. Derm of head not fused with cranium but dorsal surface of cranium exostosed. hand Color in Dorsum 12. — Dimensions. Remarks.3 mm. Indistinct black dots present on the folded skin along the sides. another dark stripe originates behind the eye. 17. There are striking differences in color. It was with hesitation that we referred this Colombian specimen to the species G. head width. skin of throat and chest smooth. 58. the last row just dorsal to the vent. 19. —A dark frog with little discernible a uniform dark brown. On direct comparison with the type of cornutum (a female).7 mm. heel reaches well past tip of snout. no traces of a skinfold across chest. Indistinct but discernible dark blotches on the upper lip. heels considerably overlap.. . Dorsal skin above lumbar region very flat medially. Thigh a lighter pattern. vocal sac not apparent. with raised ridges of skin along each side forming an incipient pouch.. knee extends to axilla.. 37. head length.6 mm. a dark brown stripe originates at tip of snout. passes upward between the nares. in having well-developed trace of web between third and fourth toes (absent in the type).2 mm. passes posteriorly above the tympanum and along the side to the lateral margin of the egg sac opening. in not having the parietal marginal walls nearly so elevated. . Skin of upper parts smooth. the first of these rows is between the eyes. this specimen (a male) differs as follows: in its smaller size (47.2 mm. the color fades into the ventral ground — . tibia.5 mm. Head and body. intersquamosal shelf incomplete and not forming a straight posterior margin at the back of the skull.5 mm. as is characteristic of male Gastrotheca. when hind legs are bent at right angles to body. foot. belly and lower femur uniformly granular. for the type). There is a spine of skin forming a supraorbital spine projecting outward from each upper lid. femur. No patagium. There are eight series of tiny.2 mm. brown with four narrow dark brown crossbands. The type has a small but very distinct dermal heel appendage. although this may be sexual dimorphism.

Peters. It lacks superciliary processes and dermal heel appendages. p. Pensilvania). Canthus rostralis sharply defined. ECUADOR: PANAMA: Cachabe\ Pacific side. transverse series lying close together between the small. above the vent. separated from each other by an interval equal to about two-thirds their distance from eye. —MZUM 73242 (holotype). USNM Gastrotheca nicefori Gaige Figure 1933.S. sloping in profile. separating dorsal yellow from gray color of posterior thighs. 15. but these are not definite warts as they are in the Colombian specimen. transverse line distinct species. — A large Gastrotheca with the derm of the head cowith the roof of the skull and with the intersquamosal shelf complete and forming a nearly straight transverse margin at the back of the head. p. Tacarcuma village. The described specimen also was compared directly with the type of angustifrons.—J.49] (type). From medemi it can be distiguished by its broader head and its longer leg (tibia greater than half the distance from snout to Diagnosis.162 [1947.third that from eye.28.4. 1963.2. There are two bright yellow spots below the vent. 1944b. tongue two-thirds as wide as mouth opening.500 m. Colombia. It is somewhat similar to that species in color. NATIONAL MUSEUM BULLETIN 288 There is a distinct. CNHM 54718. which is dirty yellow with darker mottlings. It may well be that after more information becomes available on cornutum our specimen will be found to represent a color. Vomerine teeth in two short. the upper jaw not extending considerably beyond lower. broadly rounded. p. pale. until then. 21. Caldas. p. 141795. Description. 11 Gastrotheca nicefori Gaige. rounded choanae. 1 (type locality. ossified tip of urostyle). Darien. loreal region slightly . somewhat U-shaped when viewed from above.— Gorham. nostrils more lateral than superior. BM 98. its posterior border slightly free and shallowly but distinctly notched. from Pensilvania.— 170 U. 1. This species can be immediately distinguished from all Colombian Gastrotheca except medemi by having the cranial derm fused to the skull and by having the straight posterior margin to the intersquamosal shelf. 1952. Dunn. rather heavy.22. their distance from end of snout about one. Snout moderate.. but angustifrons lacks the superciliary processes and really seems to be quite distinct. maneat! Specimens Examined COLOMBIA Cauca: Rio San Joaquin. not projecting. Caldas. 405.

the web on fourth toe reaching the proximal one-third of the antepenultimate phalanx. separated from eye by a distance nearly equal to two-thirds its own diameter. no projecting rudiment of a pollex. CNHM 6388G (same size). No patagium. when hind legs are bent at right angles to body. in postaxillary region a little broader than greatest width of head. about three-fifths the diameter of eye. heel reaches anterior margin of eye. fourth finger considerably longer than second. no dermal appendage on heel. Body not elongate. Eye moderate. not particularly prominent. when hind leg is adpressed. heels overlap greatly. its diameter equal to its distance from nostril. Tympanum very distinct. which slightly more than covers the tympanic area. disk of fourth just covering the tympanic area. which is slightly greater than distance between nostrils. a rather narrow. knee and elbow meet. Gastrotheca nicefori. Toes about one-half webbed. Skin of upper parts smooth. glandular bony ridge passes above Figure 11.— FROGS OF COLOMBIA — COCHRAN AND GOIN 171 concave and but slightly oblique. interorbital distance about twice the width of upper eyelid. reaching to middle of disk of third. fifth toe about a disk longer than third. Fingers with a trace of web at base. . when limbs are laid along the side. the upper lip not flaring out strongly below it. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. no tarsal ridge.

belly — — A light taimish brown above. Meta.3 mm. vocal sac not apparent.4 mm. Pico Renjifo. intersquamosal shelf complete and forming a nearly straight posterior margin to the skull. Specimens Examined COLOMBIA Antioquia: Yarumal. Most of the specimens are rather uniform brown. Serrania de Diagnosis. narrow stripe posteriorly. Colombia.. Skin of head co-ossified with skull. whereas a large female has a head-body length of 82.0±mm.. —MZUM 123068. for the type measures only 66 millimeters and another adult female at hand is only 67 millimeters in length. nearly straight margin at the back of the head and with the cranial derm co-ossified with the roof of the skull . Medem. —CNHM 81367. slightly darker Color in alcohol. we tentatively assign it on the basis of geography and its well-developed intersquamosal shelf. CNHM 63886. which are rather dusky. 92265. 25. Cundinamarca: Fusagasuga. 1. from Yarumal. roof of skull exostosed. It is with some hesitation that we include USNM 152053. Since its smaller size (head-body 50. below.0 millimeters. head width. foot. 172 U. MLS 208. USNM 152053. MLS 343. Meta. Gastrotheca medemi. MZUM 73242 (holotype). MLS 343. —A large Gastrotheca with the intersquamosal Paratype. shelf forming a complete. NATIONAL MUSEUM BULLETIN 2 88 skin of throat and chest finely granular. head length. 46. hand 18. femur. a typical adult Variation.. 400 meters. Serrania de La Macarena. — male having a head-body length of 61.3 mm.500 meters. Caldas: Pensilvania. particularly the underside of thighs... new species Figure 12 Holotype. from Sutatenza. but one adult female. the tympanum. 71218 (paratype).4 mm.) may indicate immaturity and since we know rothing of ontogenetic changes in this species. That specimen has a pattern on the dorsum of scattered dark spots and the head is narrower than typical for nicejori.7 mm. collected by F. Dimensions.. that of and lower femur uniformly and coarsely granular. Colombia. here. Boyaca: Sutatenza. 31. an adult female from Cano Guapayita.3 millimeters. The male is smaller than the female. adult male.S. La Macarena.2 mm. All mature females are not this large however.0 mm. 36. is quite dark along the sides and has the dark lateral color separated from the brownish dorsal color by a distinct. no inguinal gland. 23.. a welldeveloped skinfold across chest. Head and body 65. tibia.

nicefori. and smaller tympanum. nostrils more lateral than superior. broadly rounded.— FROGS OF COLOMBIA Gastrotheca from which medemi probably it differs is — COCHRAN most AND GOIN 173 closely related to G. rounded lying between the small. their distance from end of snout about two-fifths that from eye. Vomerine teeth in two short. — Description of holotype. shorter legs. its posterior border free and shallowly notched. Gastrotheca medemi. rounded choanae. somewhat U-shaped when viewed from above. modas slightly Figure 12. the upper lip flaring out below it. paratype. separated from each other by an interval equal to about three-fifths their distance from eye. Eye moderate. Snout moderate. the upper jaw extending a little beyond lower. Canthus rostralis well defined. in its narrower head. loreal region concave and somewhat oblique. tongue four-fifths series. wide as mouth-opening. somewhat rounded in profile for a Gastrotheca. CNHM 81367 (same size) . very slightly projecting.

greater than width of upper eyelid. palpebral membrane not reticulate. Tympanum very diameter of eye. when limbs are laid along the side. 29.7 mm. 23. between axilla flecks in the mm. in postaxillary region a little narrower than greatest width of head. tibia. Remarks. Top of head and dorsum are pale brown without any discrete pattern.5. separated from eye by a distance equal to its own diameter. 43.5.8 Color in alcohol. 29. bordered above by a bony shelf.. — Dimensions. disk of fourth toe just covering the tympanic area. its diameter equal to its distance from nostril. 74. fourth ringer slightly more than a disk longer than second. Skin of upper parts rather smooth. the pale flecks being better developed but by no means conspicuous. heels meet snugly. Fingers unwebbed. no between projecting rudiment of a pollex. head length. A distinct nuchal dermal fold. 62.4. reaching to disk of third which slightly more than covers the tympanic area.2 mm. Body not elongate. heel-to-toe.1 mm.174 U.8. about one-half the is nostrils. Head and body. that of belly and lower surface of thigh uniformly and very finely granular. Skin of head co-ossified with skull. Adult male. femur.. No patagium. 21. 25. 33. tibia.3 mm. A rather narrow glandular ridge supported by a bony shelf passing above upper part of tympanum. cadaverish brown below. 25. a distinct oval inner and a smaller. 54. interorbital which distance slightly- about equal to distance distinct. no ulnar ridge. third and fifth toes subequal. roof of skull strongly exostosed. head length. — The paratype is a large female with the following millimeter measurements: head and body.2.7 mm. hand. Toes slightly more than one-third webbed. no tarsal ridge.. a skinfold across chest. Side and groin moderate brown with a few. 37. head width. —Pale brown above. Anterior and posterior faces of thighs similar in color to the side. 22... A frontoparietal-squamosal arch which is complete to the rear of the skull. heel-to-toe. knee and elbow fail to meet. femur. no dermal appendage on heel.S. 27.5. Posterior surfaces of thighs are dark brown with mottlings . pale just anterior to groin. NATIONAL MUSEUM BULLETIN 288 erately prominent. the web on fourth toe just reaching the base of the antepenultimate phalanx. brown area indistinct.4 mm. heel reaches almost to anterior margin of eye. vocal sac apparently internal. hand. head width. On each side behind the tympanic region is an area of dark brown that breaks up posteriorly into a series of dark mottlings along the side.. when hind legs are bent at right angles to body. indistinct rounded outer metatarsal tubercle. Underside of chin and throat a little darker than belly. skin of throat and chest smooth. when hind leg is adpressed.2. An incipient pouch on the posterior portion of the dorsum.

Norte de Santander. p. and without superciliary processes. A Gastrotheca with the cranial derm free from the head. Gastrotheca helenae. This species can be distinguished from all other Colombian Gastrotheca except mertensi and aureomaculata by the above combination of characters. Dunn. From those two species. Gastrotheca helenae — — Diagnosis.— FROGS OF COLOMBIA — COCHRAN AND GOIN 175 of light tan. . without an intersquamosal shelf forming a nearly straight margin at the back of the skull. holotype. Paramo de Tama). Gastrotheca lielenae Dunn Figure 13 1944. p. 1963. 1944b. without dermal appendages on the heels. Gorham. p. 404 (type locality. it can be distinguished at a glance by its distinctive pattern. 21. 44. MLS 268 (same size). Exposed ventral surfaces are brownish gray with the underside of chin being a little darker than the belly. 1957. and indeed from all Gastrotheca known to us. Colombia. Figure 13.

rounded when viewed from above. loreal region nearly flat and nearly vertical. not projecting. skin of throat. Snout short. nostrils more anterior than superior or lateral. when hind legs are bent at right angles to body. NATIONAL MUSEUM BULLETIN 288 — MLS 268 (holotype). then posteriorly through the lower margin of the upper eyelid. Canthus rostralis well defined. which is relatively narrow. chest. Tympanum distinct. when limbs are laid along the side. —A handsome frog with a dorsal ground color of cream with the following dark brown dorsal pattern: a fine line from the tip of the snout passing dorsally and forking to pass through each nostril. its diameter equal to its distance from nostril. 20. 33 mm. its posterior border free and with a short but acute notch. disk of fourth toe about covering the tympanic area.7 mm. knee and elbow overlap slightly. third and fifth toes equal. a weakly developed dermal appendage on heel. . no traces of a skinfold across the chest.2 mm. tibia. slit-like opening. 48... when hind leg is adpressed. femur. metatarsal tubercle not distinct. from Paramo de Tama. 25. about one-third the diameter of eye. separated from eye by a distance nearly equal to twice its own diameter. Toes slightly less than one-half webbed. foot. reaching to disk of third which covers the tympanic area. interorbital distance slightly more than twice the width of upper eyelid.S. thence posteriorly above the tympanum to the shoulder region where . separated from each other by an interval equal to about one-third their distance from eye.5 mm. heels overlap distinctly.5 mm. Dimensions. Body not elongate. belly. Norte de Santander. in postaxillary region much broader than greatest width of head. the upper jaw not extending appreciably beyond lower. and lower femur uniformly and very finely granular. hand 21. fourth finger considerably longer than second. No patagium. No narrow glandular ridge encircling upper part of tympanum. . not prominent. bony intersquamosal plate forming a straight margin at the back of the skull. Fingers entirely free. skin of head not co-ossified with skull but roof of skull exostosed.. 35. Colombia. but slightly greater than distance between nostrils. 62. rounded.2 mm. head length. rounded choanae. Vomerine teeth in two small. a rounded but not spined rudiment of a pollex. Head and body. 1 mm. tongue three-fourths as wide as mouth opening. heel reaches to middle of eye. no vocal sac apparent. — . without a flat. Color in alcohol. Eye small. nearly truncate in profile but sloping slightly posteriorly from the vertical. their distance from end of snout about one-sixth that from eye. no tarsal ridge. oval series lying on a level with the posterior borders of the small. head width. the upper lip not flaring out strongly below it. Description. Skin of upper parts highly smooth. web extending to middle of antepenultimate phalanx of fourth. 176 U. A dorsal pouch present which opens posteriorly by a vertical.

this new species can be differentiated from bufona and cornutum by the absence of superciliary processes. Niceforo Maria. about 20 millimeters in head-body length. from nicefori and medemi by having the cranial derm free from the skull. Color in life. In life the ground color is inky blue but greenish gold spots are so thickly spaced that the frog so that from a heels. an adult male. and from helenae. another pair on each upper eyelid. collected at Moscopan. Two gray-green (in life) young. —-A Gastrotheca without the cranial derm co-ossified with the skull. and continue posteriorly along the midline of the back as a broadening line that posteriorly broadens to completely encompass the opening of the pouch. There is a fine dark line along the entire margin of the upper lip. by Marte Latham. at San Agustin. — Remarks. straight margin at the back of the head. —CJG 2307. Although Dunn (1944b. A few dark stripes cross. the top of the thigh. mertensi. underside of the limbs. there is a band of brown across each wrist. We can offer no suggestions as to the relationship of helenae. 404) described this species as having the derm of the head fused with the skull. collected in Huila. a small village about 25 kilometers west of Holotype. both above and below. p. bia. In preservative it is black with numerous small spots of yellowish white. Diagnosis. new species Figures 14-16 — CNHM 69701." They show it of fine lines originate — — the same pattern as the parent. ColomSan Antonio. it little distance appears bright green. There are a few spots on each side of the middle of the back. Colombia. without dermal appendages on the and with the intersquamosal shelf incomplete posteriorly does not form a complete. the head is really quite free. Cauca. "Bright yellow with reddish brown and white marks. The entire venter. otherwise the tops of the limbs are cream with scattered dark spots. after which it abruptly terminates. and monticola argenteovirens by having the dorsum and venter thickly covered with light spots. The specimens in the series MLS 268 (the type and two juveniles) were the only ones examined. From the other species of Gastrotheca now known from Colombia. and the sides of the body are a rich chocolate brown. transverse. Note on life history. pass medially and posteriorly to the middle of the head where they fuse. at 2. or nearly cross.300 meters. were "born after capture. without superciliary processes.FROGS OF COLOMBIA — COCHRAN AND GOIN 177 merges with the dark brown ventral and lateral color. . Gastrotheca aureomaculata." according to original field notes shown to us by Hno. Paratype.

2 "3 m 1:1 I s II a M O 13 . NATIONAL MUSEUM BULLETIN 288 o Qi .S. &S o .178 U.

Skinfold across chest a heavy pectoral fold. heels overlap patagium. vocal sac not pronounced. the web on fourth toe reaching the distal end of the preantepenultimate phalanx. that of belly and lower surface of thigh uniformly more distinctly so. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. no ulnar ridge. the upper jaw extending very little beyond lower.5 mm. head width. when limbs are laid along the side. The of the proximal two-thirds of the thigh to them and to those on the granules on the undersurface finer than those adjacent much belly. heel reaches posterior margin of eye.. disk of fourth toe just covering the tympanic area. — . head length.5 mm. 22. tongue two-thirds as wide as mouth opening.. rounded when viewed from above. Skin of head not co-ossified with skull. Snout not large. rounded choanae. lying close together on a level with the posterior borders of the small. In preservative this is a black frog with extremely numerous small yellowish white spots about one millimeter .. thus forming a rather distinctive patch. Body not elongate. heel-to-toe. no projecting rudiment of a poll ex. truncate but slightly sloping in profile.3 mm. palpebral membrane not reticulate. A fairly skin of throat and chest finely granular. its posterior border and very shallowly notched. Skin of upper parts finely granular. the upper lip flaring out but slightly below it.4 mm. reaching beyond middle of disk of third which covers the tympanic area. their distance from end of snout about one-third that from eye. adult male. femur. — Dimensions. Head and body 66. no tarsal ridge. somewhat transverse —COCHRAN —Vomerine AND GOIN 179 teeth in two moderately large series. tibia. separated from each other by an interval equal to about two-thirds their distance from eye.5 mm. 32. Toes less than one-half webbed. Tympanum very distinct. Eye neither large nor prominent. third and fifth toes subequal. its diameter equal to its distance from nostril. Color in alcohol. fourth finger considerably longer than second.5 mm. 25. roof of skull strongly exostosed. Canthus rostralis well defined. Fingers with but a trace of web at base. interorbital distance one-third greater than width of upper eyelid. An incipient No legs are bent at right angles to body. about two-thirds the diameter of eye. separated from eye by a distance nearly equal to its own diameter. in postaxillary region nearly equal to greatest width of head. which is in turn about equal to the distance between nostrils.3 mm. distinct glandular ridge encircling upper part of tympanum . considerably projecting. hand. 50... no dermal appendage on heel. broadly cordiform. 25. marsupium discernible in supra-urostylic region. knee and elbow overlap slightly free slightly. when hind appreciably. nostrils more lateral than superior. when hind leg is adpressed.FROGS OF COLOMBIA Description of holotype. loreal region flat and nearly vertical. 36.

each of which has three weakly developed toe buds. spots along side larger and more sparsely spaced so that the ground color is more evident along the side. these spots larger than those iris — metamorphosed until May 17 May when 15. The first tadpole tions of white. — Tadpole. There are two complete rows of teeth dorsally and below the mouth are three rows. belly surface silverish gray with thickly set spots and vermicula- in diameter scattered over — on the dorsal surface. (See Fig.2 millimeters. On the belly the spots tend to run together to form vermicnlations. Underside of chin and throat bluish green with light and dark specks giving a salt-and-pepper effect. Concealed surfaces of thighs and axillary regions inky blue with spots more whitish than they are along side of body.6 millimeters is head and of — body. 16. As soon as the food was put into the aquarium with them. pouch when she arrived in Gainesville. thus resembling pattern along sides. top of head and entire back thickly beset with small spots and vermiculatelike marks of a rich greenish gold. Gaslrotheca aureomaculata (X 3). NATIONAL MUSEUM BULLETIN 2 88 it. The tadpoles continued to metamorphose the steam heat was inadvertently turned on in the building over the week end.180 U.) The mouth is bordered by fleshy. above and below. One of the newly "born" tadpoles has a total length of 25. Undersurfaces of hands and feet bluish gray. Dorsal surfaces of arms and legs with the inky blue ground color of the back but with greenish gold spots larger and more sparsely spaced. the bottom two of which are complete. Hind limbs are present. or spinach were all eaten with equal enthusiasm. Dorsal ground color inky blue. Color of paratype in life. sliced cantaloup. Pieces meat or gristle. as they all died. the uppermost being interrupted in the middle.S.) Figuke 15. of which 9. The live female paratype was carrying eggs in her Life history. too high for the tadpoles The tadpoles fed avidly on nearly everything offered them. Description of tadpole. The spiracle is on the left side of the body about half-way between the head and the base of the tail. . 15. bronzy gold. tympanum bright yellow. finger-like projections all around except for about the median third of the dorsal lip. Presumably the temperature became and young. (See Fig. they would gather round it and start eating at any exposed place not preoccupied by other tadpoles. Florida. The night of April 5 she "gave birth" to approximately 70 tadpoles.

The young were bright green life. April-May 1957. In addition to the type and paratype. without superciliary processes. this female was collected at 10 to 15 miles west of Playa de Oro. an adult female from Moscapan. Colombia.both are really from the same vicinity. — 337-262r-70 13 . Chocd. as the two localities are in the same mountain range only about 40 kilo- meters apart. all from Moscop&n. Colombia. CJG 2305-6 (alive). AND GOIN is 19.—MZVM 121025-30. collected by Marte Latham.4 millimeters in head and heavily pigmented but no discernible pattern can in the preserved specimen. a dried. Colombia. Diagnosis. Gastrotheca mertensi. A Gastrotheca without dermal appendages on heels. The female in life is a waxy green on the exposed dorsal surfaces and dark inky blue on the concealed surfaces of the thighs and under the arms. and with the cranial derm free from the cranium. 121024. faded female (USNM 152333) received from the National Zoological Park seems to be referable to this species. Cauca. According to the data that accompanied the specimen. It made out is 181 of tadpole. Gastrotheca aureomaculata. Paratypes. recently transformed individual body be 16. An intersquamosal shelf is present but incomplete laterally. — Remarks. Sometimes the female has the top of the head change to a pale blue. new species Figure —MZUM 17 Holotype.—COCHRAN FROGS OF COLOMBIA Figure A in — Mouthparts length. by Marte Latham. USNM 148567-8 (alive). In life the male ranges from bronzy to greenish on all exposed dorsal surfaces with a dark. Cauca. inky blue color on the concealed surfaces of the thighs and under the arms. 152334-74 (41 young). Although the holotype came from the state of Huila and the paratype from the state of Cauca. In preservative adults of both sexes are gun metal blue above and dirty gray below. thus not forming a complete transverse shelf at the rear of the head. 121031 (7 young).

— 182 U. — Figure 17. . tongue half as wide as mouth opening. NATIONAL MUSEUM BULLETIN 2 88 all Colombian Gastrotheca exand aureomaculata by the absence of both superciliary processes and dermal appendages on the heels and by its having the cranial derm free. Snout large. the upper jaw extending but slightly G. transverse series. Its uniform pattern easily distinguishes it from both helenae and aureomaculata. heavy. its posterior border free and distinctly notched. somewhat elongate and triangular for a Gastrotheca when viewed from above. rounded choanae. truncate and slightly sloping in profile. mertensi can be distinguished from cept helenae. From monticola argenteovirens it differs in its larger size and in the fact that the adults are uniform below rather than mottled. longer than wide. MZUM 121024 (same size). Description ojholotype. Vomerine teeth in two short.S. monticola argenteovirens . Gastrotheca mertensi. lying close together between the small. holotype.

but distorted by having the theca packed with eggs. The females are ticola agenteovirens. not projecting. Eye neither large nor prominent. when limbs are laid along the side.. reaching to middle of disk of third which covers about four-fifths the tympanic area. the web on fourth toe reaching the base of the antepenultimate phalanx.5 mm.2 mm. no dermal appendage on heel. monserved series of adults. no ulnar ridge. Skin of head not co-ossified with skull. to bluish —There is gray on little all variation all dorsal surfaces. — Dimensions. about three-fourths the diameter of eye. When hind leg is adpressed.. No patagium. Color in alcohol.— Gun-metal blue on from dirty gray apparent. No pattern apparent among the pre- somewhat larger (two adult females. and 62. Body somewhat elongate. adult female. heels overlap greatly. 39. which is relatively wide and slightly greater than distance between nostrils. heel-to-toe.8 and 71 mm. tympanum. Skin of upper parts smooth but with tile-like pattern due to the eggs in the theca. a distinct oval inner and a smaller but distinct rounded outer metatarsal tubercle. 28. hand. top of forearm. head length. no projecting rudiment of a poll ex. dorsum. roof of beyond lower. Tympanum very distinct. from end of snout about two-thirds that from eye. Can thus rostralis well defined. Head and body.. separated from each other by an interval equal to about two-thirds their distance from eye. femur. skin of throat and chest. 64. 58 mm. shank.1. when hind legs are bent at right angles to body. thus not forming a complete transverse shelf at the rear of the head. loreal region concave and nearly vertical. 35.) and one of the males has a more mottled belly surface. Intersquamosal shelf incomplete laterally. interorbital distance slightly greater than width of upper eyelid. hand. its diameter equal to its distance from nostril. and lower surface of thigh uniformly and very finely granular. in head and body length. knee and elbow slightly overlap. tibia. their distance skull strongly exostosed.2 mm. third and fifth subequal. no vocal sac. and foot. 25.7. A rather narrow glandular ridge passes above tympanum. 73. three adult males.3 mm. 64. 28.. — Color in life. head width. no tarsal ridge. palpebral membrane not reticulate. Toes about one-third webbed.1 mm. Variation. Male: Very pale bronzy green on top of head.8 mm. belly. in this respect approaching the appearance of G. Fingers with just a trace of web at base. heel reaches anterior margin of eye. disk of fourth covering about two-thirds the tympanic area. top of upper .2 mm. separated from eye by a distance nearly equal to its own diameter.. traces of a skinfold across chest.. ranging ventral surfaces.FROGS OF COLOMBIA — COCHRAN AND GOIN 183 nostrils more lateral than superior. 73. fourth considerably longer than second.

A living male (in the office of the junior author) calls Voice. As the above data indicate. No pattern was discernible in the time of calling. When hatched the tadpoles had a head-body length of about tail length of about 15 millimeters. the largest individual had a head-body length of 30. underside of chin. April 8. or at night. Remarks. We believe that G. When the last specimens of this series were preserved on August 31. snarl is — Florida. Two pairs of specimens were sent alive to the senior Life history. author in March 1962. the tadpoles do not all grow at the same rate. The body was about 5 millimeters in diameter. On April 12 a piece of pork chop (cooked) was put in the aquarium and they immediately gathered first 10 millimeters and a around it and started feeding on it like pigs at a trough. and along sides of body inky blue. single notes.S. mertensi probably is most closely related to marsupiata. arms. concealed surfaces of thighs. 184 NATIONAL MUSEUM BULLETIN 2 88 arm. The call is made up of two gray.S. and the top of the head tends toward sky blue rather than bronze. The gravid female at the Museum give birth to 96 tadpoles on the weekend of April 6-8. the tympanum and iris bronze. and belly bluish is some metachrosis. and iris bright golden bronze. irregularly during the day and night. and the explosive notes which follow are very similar to a single note of Rana virgatipes. The egg capsules from which the tadpoles emerged were found in the water and measured. National Museum for observation and the other pair was sent to the junior author in Gainesville.U. The gravid female and a male were retained at the U. While still in the water they ranged from four to five millimeters in diameter. but no noticeable bronze along the sides. explosive. as the dorsal ground color varies from a greenish bronze at times to a clear pale bronze at others. as in the male. 43 were kept at the Museum for observation and 40 were sent to the National Zoological Park. but since we do not have first-hand knowledge . Female: Similar in pattern to the male but the dorsal ground color is a waxy pea green with no trace of bronze. although they are not given nearly so rapidly. Thirteen of the tadpoles were found dead on Monday morning. the last one transformed on August 24. Of those remaining. metamorphosis occurring in this one lot from about mid- May to August — 24. The specimens at the Museum started metamorphosing in May. since he would call in morning. by Marte Latham. in afternoon. to eight short.4 millimeters. and there — parts — first a series of six to eight snarls followed The by a series of four not too unlike that of Scaphiopus. 1962. throat. On April 9 the tadpoles were observed feeding on Spirogyra in the aquarium. anterior portion of side below the dorsolateral stripe.

rounded choanae.000 meters. separated from each other by an interval equal to about two-thirds their distance from eye. — Diagnosis. However. Specimens Examined COLOMBIA Cauca: CJG Moscopan. — 1963. All the specimens we have seen have come from Moscopan. and both he and his colleague. Moscopan. Eye moderate. p. Gastrotheca monticola argenteovirens (Boettger) Figure 18 1892. is not a town or city but a region near the paramos. 46 (type locality. 21. Colombia. very slightly rounded in profile. Mertens. Can thus rostralis sharply defined loreal region nearly flat and nearly vertical. not particularly prominent. A small Gastrotheca without superciliary processes. Robert Mertens compared mertensi directly with the type of monticola argenteovirens. Narifio . tongue two-thirds as wide as mouth opening. Mertens. 2. Dr. nostrils more lateral than superior. it is surely most closely related to monticola argenteovirens. rounded when viewed from above. 121025-30. Klemmer. Snout short. slightly projecting. 1922. and with the intersquamosal shelf incomplete posteriorly so that it does not form a straight margin at the rear of the skull. Colombia. Dr. p. to G. MZUM USNM 121024 (holotype). p. state that the two specimens cannot be of the same species. 121031 152334-74 (41 young). It is reached by taking the road from Popayan up past Purace* and across the paramo. Vomerine teeth in two short.— Gorham. 1933. theca. This form is more similar Gastrotheca but it CNHM Description— 54811. 259. their distance from end of snout about one. from Guaitarilla. 163. mertensi than to any other Colombian can be distinguished from that species by its small size and by the fact that it tends to be mottled below rather than uniformly colored. Vellard. Hyla argenteovirens Boettger. lying between the small. 7. p. without dermal appendages on the heel. with the cranial derm free from the skull. FROGS OF COLOMBIA —COCHRAN AND GOIN 185 Andes south of Colombia we Of the Colombian species of Gastro- of all the species of Gastrotheca in the cannot be dogmatic about this. in the state of Cauca. 148567-8. rounded. the upper jaw extending a little beyond lower. — 1957. p. Popayan). Gastrotheca marsupiata argenteovirens. its posterior border slightly free and shallowly notched. 2305-6.— . the upper lip not flaring out appreciably below it. although an often-cited locality. very broadly cordiform.third that from eye. Colombia. 1933. (7 juv.). That they are specifically distinct is borne out also by the fact that both forms occur at Moscopan and retain their distinctness there. contiguous series.

Gastrotheca monticola argenteovirens: Narino. 18.— 186 Figure U. and .S. NATIONAL MUSEUM BULLETIN 2SS CNHM CNHM 54811 (at top) from 63887 from San Pedro in Antioquia (both same size).

a well- developed skinfold across chest. no projecting rudiment of a pollex. 26. that of belly and lower surface of thigh uniformly granular. Skin of co-ossified with skull.. above upper skin of throat and chest very finely granular.. separated from eye by a distance nearly equal to one-half its own diameter. This is inflated and contains eggs. roof of skidl strongly exostosed. The opening is a longitudinal front of the vent. adult female. no dermal appendage on heel.. when limbs are laid along the side. Preserved specimens from San Pedro in Antioquia are smoother skinned. no vocal sac. and are more somber in appearance. Some geographic variation that may be significant is indicated. knee and elbow fail to meet. fourth finger considerably longer than second. Toes slightly less than one-half webbed. the web on fourth toe reaching the base of the antepenultimate phalanx. showing less mottling on the belly and concealed portions of the thighs and groin than do the Nariflo and Cauca head not — — — . 59 mm. heel reaches middle of eye. Variation.FROGS OF COLOMBIA — COCHRAN AND GOIN 187 diameter slightly greater than its distance from nostril. no ulnar ridge. Tympanum very distinct. Color in alcohol. disk of fourth toe just covering the tympanic area.4 mm. 17. A well-developed dorsal marsupium present. head length. heel-to-toe.. heels overlap somewhat.4 mm. when hind legs are bent at right angles to body. part of slit. femur. 27.' In color and pattern the series is rather uniform.7 mm. Dimensions. but in general they are much the same. reaching to middle of disk of third which covers the tympanic area. 18. Some individuals are more heavily mottled below and some have more prominent postocular dark stripes. Chin and throat gray. Similar but less distinct mottlings occur on the undersurfaces of arms and legs.' Head and body. Skin of upper parts smooth.9 mm. about three-fifths the diameter of eye.. third and fifth toes subits equal. which is slightly greater than distance between nostrils. interorbital distance half again as great as width of upper eyelid.. have more prominent dorsolateral folds. palpebral membrane not reticulate. Body not elongate.3 mm. head width. when hind leg is adpressed. Fingers webbed at base. tibia. Dark lead gray on all dorsal surfaces. belly surface very light gray with thickly scattered flecks of brown. 7 millimeters long. in postaxillary region a little narrower than greatest width of head. tarsal ridge. a distinct oval inner and a smaller but distinct rounded outer metatarsal no tubercle. hand. 40.7 mm. that lies 10 millimeters in A rather narrow glandular ridge passes tympanum. No patagium. Along the side between axilla and groin and on the anterior face of the thigh neckings of the pale gray ventral color give a somewhat mottled appearance. 20. web reaching base of antepenultimate phalanx of third finger.

CNHM Quintana (Moscopan). for the late E. In the second case the difference may be due either to the fact that we here recognize groups heretofore not recognized or groups that differ either in generally are not represented in southeastern Brazil.188 U-S. expanded in all except some of the larger present. CNHM 63887-8. and 54824. a well-developed quadratojugal.000 m.- p. that the San Pedro form might be new. 32 (type species. a female) to our good friend Dr. 1955) we have divided — the large. R. In the former case. or parasphenoid. 44048-9. 78300. vocal pouch in male. no backward projecting process on ischium. San Pedro. 18). tympanum present and usually fairly distinct. MCZ 20936-7. 2676 (photograph 43981. Antioqtjia: 71217.. Dr. NariSo: Guaitarilla. larger groups. —Without teeth on mandible. . under arqenteovirens. 90599. CNHM 54723. In other cases the groups here recognized being less inclusive or in not being recognized as groups in the Brazilian work. 54844-5. Robert Mertens in Frankfurt and asked him to compare them with the type. 24916. the difference is most often due to the fact that with more material available we have been able to further subdivide some of the earlier. if palatine. Following the method used by the senior author in her Frogs oj Southeastern Brazil (Cochran. tongue fairly well fused behind. For this we — are very grateful. 54792. Hyla Laurenti. sacral diapophysis well forms. To lima: Quindfo Mountains. Since the correct assignment of the name "argenteovirens" has been a problem. unwieldy genus Hyla into several groups. 89454. Remarks. We are not the first to notice this. He wrote that both he and his colleague. 2. 56507. we sent two of the large series of specimens from Nariiio (CNHM 54823.. Dunn made a handwritten note on a manuscript key to Gastrotheca. Tuqueres. In many cases these groups coincide with those that the senior author recognized in southeastern Brazil. MCZ 16080-1. CNHM 3. Remarks. MCZ 8224-6.000 m. cranial derm not fused with skull except in a few West Indian species. a male. median and subgular. of holotype). neither eggs nor young carried on back of female. 54793-843. Popayan NMS 9757-60. CNHM Genus Hyla Laurenti 1768. Hyla viridis Laurenti). Klemmer. Diagnosis. NATIONAL MUSEUM BULLETIN 2 88 specimens (see Fig. no well-developed ventrolateral gland along each side. Cundinamarca: Bogota. had made the comparison and both were of the opinion that the two specimens were conspecific with the type. Specimens Examined COLOMBIA MZUM Cauca: MCZ Medellin. pupil horizontal.

We do not believe of species of advisable to present a key to the entire it Colombian Hyla. lanciformis group. H. lanciformis.— Goin and Layne. instead. A rather large Hyla with a pointed snout. 63. group. rubra group. squalirostris 1. 9. 1 species carri group. p. 15. 22. p. 19. 3 species 2 species LANCIFORMIS group This group consists of large tree frogs with a white margin on the upper lip. p. fingers showing a trace of web at base. maxima group. 649 Goin.— Andersson. p.— Gorham. Surely AND GOIN when 189 in Colombia. Ecuador). 1945. — Hyla — Nieden. 4 species 20. 1882. 1 species 13. 119. — Diagnosis. 3 species 6. 1 species 7. albomarginata group. Hyla microcentra Werner. indicate the relationship of our species groups to extralimital forms. p. 3 species group. buckleyi group. p. 1884a. 8 species 22. 109. — 268. boans of southeastern Brazil. crepitans group. 1 2. geographica group. leucophyllata group. 22. pointed heads. Hyla lanciformis (Cope) Plate 25a-c 1870. 4. 1 species 8. 1903. 1958. p. 3 species 5. minuta group. at least two other forms are in this group. 1923. 1958. a conspicuous white margin on the . microcephala group. palpebrogranulata 1 species 16. 2 species 17. and additional studies have been made on the frogs of other geographic areas in South America there will be some rearrangement of the groups we here recognize. 1952. 2 species phantasmagoria group. 10. p. 361. p.— Lutz and Kloss. hobbsi group. variabilis group. p. labialis group. mulitfasciata of northeastern South America and H. granosa group. 1 1 species species 21. parviceps group. we have attempted list to prepare adequate diagnoses for each of the species groups and have then presented a key to the Colombian species within each group. 1 species marmorata group. The groups we here recognize in Colombia and the number of Colombian species now known in each group are as follows: 12. Boulenger. but for the time being all we can do is to arrange the Colombian species into such natural groups as we can. and when possible. 1963. — Gorham. 180 (type locality.FROGS OF COLOMBIA —COCHRAN At present we can recognize 47 Hyla species of these are arranged into 22 groups. 556 (type locality. Colombia). 2 species 18. species 3. 1882a. 14. 3 species 1. Along with Hyla lanciformis. 1 species 11. 2 species punctata group. 637. and reduced webbing between the fingers. p. Hxjpsiboas lanciformis Cope. phlcbodes group. 1921.

Skin of throat and chest smooth. a distinct oval inner but no distinct rounded outer metatarsal tubercle. broadly cordiform. not projecting. Snout elongate. loreal region slightly concave and nearly vertical. Skin of upper parts very finely granular. Tympanum its distance from nostril. the web on fourth toe reaching distal end of the antepenultimate phalanx. reticulate. pointed in profile. vocal sac subgular and but slightly apparent. No patagium. when hind legs are bent at right angles to body. NATIONAL MUSEUM BULLETIN 2 88 upper jaw. heels overlap greatly. separated from each other by an interval equal to about two-thirds their distance from eye. horizontal glandular white line above the vent. Toes slightly more than one-half webbed. Colombia. Eye neither large nor prominent. a weak ulnar ridge. and a transverse. reaching to disk of third which covers about one-third the tympanic area. the upper jaw extending somewhat beyond lower. palpebral width about the distance between nostrils. its posterior border slightly free and shallowly notched. line — USNM Description.S. The combination of size. . and white above the vent serves at a glance to distinguish this species from any other Colombian Hyla. white border to upper jaw. Rio Orteguaza. the upper lip not flaring out appreciably below it. Vomerine teeth in two long angulate series. separated from eye by a distance equal to one-half its own diameter. about three-fifths the diameter of eye. no tarsal ridge. Can thus rostralis strongly defined. from La Providencia. lying between the very elongate choanae. no dermal appendage on heel. Skin of head not coossified with skull. disk of fourth toe covering about one-third the tympanic area. their distance from end of snout slightly more than one-third that from eye. adult male. Nostrils entirely lateral. its diameter about three-fourths membrane not of upper eyelid.190 U. tongue three-fourths as wide as mouth opening. knee reaches axilla. no traces of a skinfold across chest. roof of skull not exostosed. Caqueta. when hind leg is adpressed. when limbs are laid along the side. third and fifth toes subequal. interorbital distance greater than which in turn is very distinct. Body elongate. A rather narrow glandular ridge passing above tympanum then continuing above shoulder as a short dorsolateral fold. web just reaching base of ante- penultimate phalanx on lateral side. Fingers with a trace of web at base. in postaxillary region a little narrower than greatest width of head. fourth finger considerably longer than second. no projecting rudiment of a pollex. heel reaches well beyond tip of snout. a narrow but distinct glandular ridge passes transversely above the vent and similar but less distinct transverse ridges rim across each heel. rather pointed when viewed from above. 152036. that of belly and lower surface of thigh uniformly and finely granidar.

Three of the specimens at hand have a dark dorsal median stripe running from the tip of the snout posteriorly to above the urostyle. heel-to-toe. The webs on the feet are — dark. small but distinct white spots on the rear of the thighs. which is probably from Cafio Guapaya. 36. posterior and ventral surfaces of thigh dark and with tiny. There seems to be no doubt that lanctformis Cope of eastern Colombia. they are quite variable in pattern. shanks. and western Amazonas in Brazil. scattered.. second. Serrania de La Macarena. 56. boans: Upper lip narrowly but not conspicuously margined with white. hand. 24 mm. The posterior surface of the thigh is rather uniform on most specimens.. but on several of them an inconspicuous but distinct row of white spots can be seen. in some the dorsal ground color is practically uniform. and white blotches in the pectoral region. differing primarily in details of — pattern as follows: landformis: Upper lip broadly and conspicuously margined with white.2 mm. Peru.5 mm. In dorsal ground color they range from pale gray to nearly black. —The series at hand are rather constant in the presence of a conspicuous white border along the margin of the upper jaw. The entire side of the face from the tip of the snout to the tympanum is a dark brownish black. The ventral surfaces are a dirty brown with distinct pale spots present on the throat and breast. While crossbars are fairly conspicuous on top of the head and dorsum of most. head length. and there are four such bands across the head and eight across the back. and the eastern Venezuelan Guayana are three very closely related forms. Color in alcohol. and forearms are crossed by dark bands similar to those on the back. head width. A dark chocolate brown frog with rather distinct transverse bars of a darker hue across the top of the head and the back. In some specimens the disks of the first. 44. there are scattered. Variation. and multifasciata Giinther of Para. tarsal portion of the feet. and third fingers are distinctly capped with bright yellow. Head and body. but these bands are less conspicuous on the limbs than they are on the dorsum..8 mm. 63. The thighs. Brazil. and in one specimen (FM 7r). The largest specimen at hand is a female that measures 83.2 mm. although most are medium to dark brown dorsally. rounded white spots. In most of the specimens this stripe is lacking. tibia. 19. a transverse white line above the vent. These transverse dorsal bands vary in width from one to four millimeters.FROGS OF COLOMBIA — COCHRAN AND GOIN 191 — Dimensions. Ecuador.. the Guianas. femur. Otherwise. Remarks. boans Latreille of southeastern Brazil.8 mm.2 millimeters in head and body length.. scattered white spots. posterior surfaces of thighs with distinct.. those on the back being wider than those on the head. 22 mm. .

151995-9. CNHM 69738. When Rivero (1961) studied the Venezuelan forms he felt that boans and multifasciata were close enough to justify calling them races of a single species. Tamboril. Montalvo. by the absence of a dermal heel appendage and of reticulations on the palpebral membranes. USNM 152318. on Rio Orteguaza. USNM 146259-62. MLS 36. 8531 (4). Bahia. UF 8528 (3). 192 multifasciata: Upper NATIONAL MUSEUM BULLETIN 288 lip narrowly but not conspicuously margined with white. 8530 (2). EPN 2600. JAP EPN 2514. 1 (type localities. Moruna-Santiago Province. p. It is beyond the scope of this of each of these forms. Rivero. 2 km. and Areal de Conquista. pi. 1767-8. EPN 1644-5. and by having at least the two terminal phalanges of the outer two fingers free of web. and perhaps all of South America. Villavicencio. Specimens Examined COLOMBIA Amazonas: Leticia. Rio Suno. Putamayo: Rumiyaco. CREPITANS group This group has only one representative in Colombia. EPN 1343. CNHM 54783-4. It is characterized by a rather large size. 81780. 1955. Caqueta: La Providencia. 1824. 2197-8. JAP 2106. 2235.000 m. Brazil). 7r. Vaupes: Mitu. Meta: Serrania de La Macarena(?). JRT 516. . Rio Bobonaza. 2. 1. posterior surfaces of thighs without white spots. Hyla crepitans Wied. Alto Napo. but before we make a taxonomic change we prefer to see material from more Amazon Valley localities than are represented in the material presently available to us.— U. EPN 1346. FM USNM 152035-7. Boyaca: Guaicaramo. but paper to draw up detailed synonymies seems that albopunctata Spix and it oxyrhina Reinhardt and Lutken are snyonyms of boans and that Werner is a synonym of lancijormis. 525. Macuma. It may be that all three forms are simply geographic races of a single wide-ranging species. Rio Vaupes. 524-7. CNHM 60341-2.S. Rio Villano. 1825. 8529 (3).. Jiboya. in Hyla crepitans Wied Plate 25d-p 1824. EPN 1484. JAP 2245. 47. 1824. 671. 1553. Rio Mecaya. ECUADOR: Napo-Pastaza Province. Rio Pucayacu. Amazon River side. p. south of Shell Mera. Rio Pindo. fig. 520-2. 1961) in our discussion of the application of the name Hyla microcentra We maxima Lauren ti. Sucua. Santiago-Zamora Province. EPN 2500. Cabeceras del Rio Solis. give the reasons for using boans Latrielle rather than albopunctata Spix as the name for the southern form (see Cochran. EPN 1662.

224.— A. p. Diagnosis.— Gorham. 320. p. 520. Boulenger. p. 2. p. heavy angulate-shaped series. p. the upper lip flaring out only moderately below it. tongue two-thirds as wide as mouth opening.— Boulenger.— Giinther. 1953. p. Hyla indris Cope. p. generally whitish. p. Suriname).— Wagler. 1914. its posterior border entirely fused and shallowly notched. p. 1882a. p. 1919. 40.— Nic6foro Maria. p. 1867. p. 707. oblong choanae. considerably projecting. nostrils more lateral than superior. p. separated from each other by an interval equal to about two-thirds their distance from eye. p.— Cope. 1926. from. 24. p.— Peracca. p. upper Rio San Jorge. 1930. 27.—Stebbins and Hendrickson. 55. 1882. 1947a.— Goin.— Hellmich. 201. 1856.— Parker. lying close together between the rather large.— Nieden. Stejneger. 1874b. — 1923. p. 38. 1867. 352.— Peters. Guib<5. 1963.— Ruthven. 104. p. 555. 1951. 1841.— Nieden. 320. Suriname). 200 (type locality. p.— Werner. Baumann. p. 549 (type locality. Hyla doumercii Dumeril and Bibron. p. p. p. 447. 93. Snout moderate. p.— Liu.— Beebe. p. p. 1939. 103. 61169. 1931. p. 160. 207. Giinther.— Boettger. 21. 1885.— Aleman. the upper jaw extending a little beyond lower. 391. 353. 1871a. Hyla xerophilla Dumeril and Bibron. — CNHM . 1939. Hypsiboas crepitans. 1892. 1963b. 200. p. 200. p. C6rdoba. 1922. 207. 181. Lutz.— Beebe. p. p. 1923. 511. p. 1919. Lutz.— B. 1903. p. 555 (type locality. 460 Boulenger. p. p. loreal region concave but not too oblique. 1871. somewhat rounded in profile. p. Hypsiboas circumdatus Cope. 1882a. Lutz. 1830. 1892. p. 163.- — —A rather large tree frog without dermal appendages on the heels or reticulations on the palpebral membranes but with a transverse. p. p. 43.— Shreve.— Miranda.— Cochran. 1955.— Miranda-Ribeiro.— Cope. 1882a. Catival. 551 (type locality. In general appearance Hyla crepitans is more similar to a halfgrown maxima or rosenbergi than to any other Colombian frog.— FROGS OF COLOMBIA —COCHRAN AND GOIN 193 — — Burmeister. p. Brazil) 1871a. 1877. 1952. 11. 1923. 1936. 283 Baumann. 352. 1951a. 121. 1951a. p.Ribeiro. p. 174. 1912. Barbour and Loveridge. Lutz. p. 1901. p. p. 40. 1964b. 66. p. 247.— Rivero. 1964a.— B. 52. 301. — — Boulenger. 1867. 199. 1951b. Nieden. 1899. 771. Cayenne). 481. very broadly cordiform. 40. p. 278. 101. 1830. p. pp. 34. 1901. 305. Description.— 1841. B. 1893. 1935. Vomerine teeth in two short.— Boettger. 1912.— Schmidt and Inger. 1927. p. 1858. 536.— Schubart. 163. 305. 1872a. their distance from end of snout about one-third that from eye. 1935. 1929. glandular line above the vent at least the two terminal phalanges and with of the outer three fingers free of web. p. p. 304.—Schmidt. p. Canthus rostralis moderately defined. p. 1963a. 482. 1937a. p. p. 55. 1932. 1882a. p. 1961. p. 92.— Crawford. Colombia. Hyla circumdata. 1949. — 108. p. 311. p. but it may be readily distinguished from both of these by having the two terminal phalanges of the outer three fingers free of web. 353. somewhat pointed when viewed from above. 34. 1959. p. 1952. p. . p.

3 mm. of medium brown. which 88 distance in turn slightly greater than distance between nostrils. on top of the head. adult female. They are present around the nares. be little variation in Some specimens are a uniform tan to brown above. about diameter of eye. head width. Variation. The posterior surfaces of the thighs are heavily mottled with brown on a pale yellowish background. separated from eye by a distance equal to about one-third its own diameter. 21. 33. when hind legs are bent at right angles to body. femur. an inner but no outer tarsal ridge. traces of a skinfold across chest.S. no vocal sac. NATIONAL MUSEUM BULLETIN very prominent. broad bands cross the dorsal surfaces of the thighs and shanks. no dermal appendage on heel. tibia.. —While structurally there seems H. dark bands. large. the web on fourth toe reaching the base of the four-fif ths the penultimate phalanx. a low ulnar ridge. disk of fourth toe covering about one-third the tympanic area.2 mm. crepitans. — — Very faint. in others the thighs are uniform dorsally and heavily some . a moderately projecting rudiment of a pollex. web on third finger extending to middle of antepenultimate phalanx on lateral side.8 mm. skin of throat and chest finely granular. Fingers webbed at base. greater than width of upper eyelid. hand. nostril. pattern-wise the opposite is to true..5 mm. when hind leg is adpressed. heels considerably overlap. Skin of upper parts smooth. Dimensions. Head and body.8 mm. 60.8 mm. head length. third and fifth toes subequal. 20. A very light brown frog with faint dorsal blotches Color in alcohol. to base of antepenultimate phalanx on medial side. palpebral its 2 diameter equal to membrane not is its distance from interorbital reticulate. a welldeveloped narrow glandular ridge encircling upper part of tympanum. heel-to-toe. reaching to proximal one-third of disk of third finger which covers about one-half the tympanic area. 37. knee reaches nearly to axilla. These dorsal blotches are rather indefinite and not at all prominent. fourth finger longer than second. others have the dorsal surface tan heavily mottled with dark brown.. and on the back from the suprascapular to the sacral region. Body elongate.. One specimen (MLS 210) is medium dark brown dorsally heavily flecked with pencil-like dark dots. that of belly and lower surface of thigh uniformly and more coarsely granular. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Toes slightly more than one-half webbed... roof of skull not exostosed.194 Eye U. 17. Tympanum very distinct. in postaxillary region definitely narrower than greatest width of head. broad. heel reaches well beyond tip of snout.8 mm. No patagium. when limbs are laid along the side. 45. In of the specimens the hind limbs are crossed with distinct. The ventral surfaces are immaculate. Skin of head not co-ossified with skull.

20946 MCZ MCZ USNM Guajira: Arroyo de Arenas. crepitans. Mariquita. MHNP MCZ 1873. 752 (type of Hyla xerophilla). USNM USNM USNM Cundinamarca: Beltran. 54778. Rio Magdalena. MCZ 24917. NHMW MZUM MZUM Ariari. USNM 152121-2. Garagoa. CJG 2456 (7).8. 16078. Puerto Berrio. Nilo. northeast of Bogota. SocarrS. MCZ BRAZIL: 2324-5.7. Puerto Boyac&. USNM 152263-76. latter only in crepitans. south of Villa vicencio. Pernambuco. USNM 152142. upper Rio Magdalena. La Selva. USNM 117513. 20940.6 millimeters in head Remarks. MCZ 7777. Specimens Examined COLOMBIA BM Atlantico: Barranquilla. CNHM 61169-70. Callias. Sincelejo-Tolu. CJG 2451. CNHM MZUM BM CJG 2291-3. San Javier. Hyla it differs xerophilla from the silvery white heel it has a wart which is enlarged to the point that it verges on beiug a dermal appendage. Antioquia: Nechi. JRT 93. Villeta. Villanueva. Los Pendales. MZUM 78303 (4) Muzo. 1909. 2448-9. CNHM 81781-2. near Villavicencio. CNHM 61168. and on each respects: It has spots scattered over the dorsal surface. Palomino. 48302. upper Rio San Jorge. Medialuna. 1929. 24) that we have noted that some small. Minas Gerais. USNM 151982. is a and body length. Finca El Cucharo. Tierra Alta. USNM 151946-61. USNM 127863-4. Told. on way to Dagua. CNHM 61752. 34. 152181. USNM 152286-7. some of them have dusky flecks along the side of the chin and throat and a few are actually dark underneath the throat. as two minor (1949.FROGS OF COLOMBIA — COCHRAN AND GOIN 195 mottled posteriorly. CM 2489. San Pedro. MCZ 2326-8. IOC 4765.23. C6edoba: Alto de Quimari. near Tocaima. USNM 152144. Rio Frio. Tolima: Espinal. Magdalena: IOC 4686. USNM 152115. while in a few hardly any markings of any type are discernible on the thigh. USNM 152209-14. 73. Bolivar: La Raya. CJG 2323-5. MCZ 16051. —We agree with Guibe synonym of #. CJG 2455. La Concepci6n. Cienaga. CJG 2450. Palmar de Varela. The three largest specimens at hand are 73. Miramonte.1. ANSP 19770. Tibii (Rio Catatumbo). Fundaci6n. 152611-2. Rio Buritaca. USNM 146264. Boyaca: Guaicaramo.11. . Bogota. 56506. p. near Medellm. 152182-3. 46908-21. Arrias do MCZ 2825. CM 7932-4. Honda. Bonita. Norte de Santander: Astillero. Sutatenza. MLS 210. La Mesa. CNHM 61751.7.7.2. 54780-2. MCZ 15062-4. Fresh material from Surinam indicates that Hyla indris should also be included in the synonymy of H. and 74. Meta: Grenada. FRENCH GUIANA: Cayenne. CJG 2315-6. Santander: San Gil. Santa Marta. USNM 151982. 2330-1. 9727. Aracataca. on Rio Villavicencio. . MLS 285. Monte Redondo. 152185. upper Rio Sinu. While the majority of the specimens are uniform pale underneath. MZUM MCZ 54619. CJG 2320-2. CNHM 81821. Catival. NHMW 82 (2). Valle: Near Bitaco.

Recito. a triangular dermal appendage on heel. 1927. 68. 188. MCZ Paria Peninsula. p. U-shaped when viewed from above. p. 1946. 1925. lying close together between the large elongate choanae (in this specimen the posterior ramus of the right vomerine patch is missing) tongue threefourths as wide as mouth opening.196 U. 1926. Esmeraldas. CJG 2402-7. 1935. from upper Rio Sinu. Cachabi. H.— Netting. 10. 3. with moderately webbed fingers.— Nieden. 1925. A group 88 17971. 1927. Palpebral membranes reticulate. 409. webs a2 . 104— Breder. 25978-3. the upper jaw extending but little beyond lower. 1898a. with or without reticulations on the palpebral membranes or dermal heel appendages. maxima.— Thornton. nostrils more lateral than superior. Las Sabanas. rosenbergi feet clear Hyla rosenbergi Boulenger Plate 26d-f 1898. broadly cordiform. 1964. nearly truncate in profile. p. maxima webbing heel. This species can be distinguished from H. C6rdoba. Canal Zone. p. p. p. . faber it differs in having the fingers fully webbed rather than having the webbing reduced. 61167. 329. MCZ SURINAM: Zanderij. their distance from end of Diagnosis. TRINIDAD: MCZ 2204. The two species known from Colombia may be distinguished by to fully the following key: a 1 . Acosta District: 26142. 264. 97. p. Description. Young specimens of maxima might be confused with H. 73. pp. VENEZUELA: Falcon. Panji. maxima by the absence of dermal heel appendages and reticulations on the palpebral membranes and by having the webs between the fingers and toes pale rather than dusky. Socarre. Vomerine teeth in two heavy /"\ series. 23024-8. 1935. 2 MCZ 17582-3. From H. of Palpebral hands and feet membrane not on hands and dusky reticulate. NATIONAL MUSEUM BULLETIN PANAMA: Summit. but maxima likewise differs from crepitans in having the fingers nearly fully webbed. MCZ 2525. its posterior border very slightly free and truncate but unnotched.S. allied to — CNHM . no dermal appendage on H. crepitans. 35. considerably projecting. Hyla rosenbergi Boulenger. p.— Liu. Ecuador).— Noble. 15. p. Snout large. Merida. In addition to the species recorded from Colombia {maxima and rosenbergi) such forms as faber and partialis are members of this group. Colombia. Guiria. MCZ MCZ maxima group of large frogs generally brownish in color. 1931. — A large Hyla with fingers rather fully webbed. 123 (type locality. closely H. p.

which is relatively wide and slightly greater than distance between nostrils. hand. in postaxillary region a little narrower than greatest width of head. roof of skull not exostosed. 44. separated from eye by a distance nearly equal to one-third its own diameter. third and fifth toes subequal. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle.. loreal region concave and very oblique. Skin of upper parts smooth.2 mm. separated from each other by an interval equal to about three-fifths their distance from eye. adult male. Fingers webbed. Head and body. a glandular ridge encircling upper part of tympanum. There are faint indications of and some more definite blotches on each side anterior to the groin but these are not arranged into distinct vertical bars although they show a tendency toward such an arrangement. 27. femur. about the diameter of eye. heel-to-toe. The webbing on hands and feet is no sacral dorsal blotches darker than the rest of the appendages. a low but distinct tarsal ridge. Tympanum very distinct. 82 mm. 337-262—70 14 .5 mm.5 mm. Body elongate. 26. but not particularly prominent. Can thus rostralis well defined.6 mm. palpebral membrane not reticulate. A dark horizontal line about a half -inch long passes through the vent and similar horizontal lines pass across each heel and become contiguous with the line on vent when legs are flexed. — way between hump and vent. vocal sac hardly apparent. 24. the upper lip flaring out strongly below it. no dermal appendage on heel. tibia. knee and elbow overlap. when hind legs are bent at right angles to body. no traces of a skinfold across chest. when hind leg is adpressed. disk of fourth toe covering about one-fourth the tympanic area. Toes nearly completely webbed. web extending to middle of penultimate phalanx of third finger on lateral side and base of penultimate phalanx on medial side. a projecting rudiment of a pollex.. interorbital distance about equal width of upper eyelid.. a tannish brown below.FROGS OF COLOMBIA —COCHRAN AND GOIN 197 snout equal to about one-fourth that from eye. that of belly and lower surface of thigh uniformly and not so finely granular. head length. Skin of head not coossified with skull. A dark chocolate brown thin line originates on top of snout just posterior to nostrils and passes backward as a median line to a point about half- — .. fourth finger considerably longer than second.5 mm. A very light brown above. its diameter not quite equal to its distance from nostril. head width. heel reaches beyond tip of snout. heels overlap greatly.7 mm. Dimensions. reaching to base of disk of third which covers about one-third the tympanic area. skin of throat and chest finely granular. Eye large. a very low and indistinct ulnar ridge. the web on fourth toe reaching to middle of penultimate phalanx and thence as a narrow margin to base of disk. No patagium. when limbs are laid along the side. 49. Color in alcohol. 60..

Character . NATIONAL MUSEUM BULLETIN —Structurally.198 U. Some are quite pale while others are quite dark. In terms of color and pattern the specimens show more variability. maxima and at first glance seems very similar to it. Remarks. rosenbergi is certainly closely related to H. All specimens show some degree on the back but the amount of contrast between the darker mottling and the lighter ground color ranges from almost imperceptible to conspicuous. The characteristics indicated in the following tabulation should help in distinguishing these two species. the differences beof mottling — tween the two seem significant and constant. but this does not seem to have any geographic significance for our palest and darkest specimens came from the same locality. Variation. Here again there seems to be no geographic correlation for the two most conspicuously mottled individuals before us came from the Panama Canal Zone and the Rio Calima in Valle. Colombia. While H. there is not much 2 88 variation in the series at hand.S.

BM 98. zebra . 1789. America).— Parker. p. 123 (type locality. 200. 1926.— Crawford. Hyla wavrini Parker. p. p. — Diagnosis. JAP CM CM Hyla maxima (Laurenti) Plate 26a-c Rana boans Linnaeus. A large Hyla with reticulations on the palpebral membranes.— Duinenl and Bibron. Taracud.— Goin and Layne.4. p. 1838.28. p.— Andersson. p. from pardalis by having reticulations on the palpebral membrane and from faber and rosenbergi by having reticulations on the palpebral membrane. 120. 1952.— Dowling. Hyla palmata Daudin.—Peracca. p. 5. Upper Orinoco. p. dermal appendages on the heels. — 1900. vicinity of Cachabi. maxima differs from geographica by having the fingers fully webbed and by the webbing being dusky to dark gray rather than clear. Rana maxima Laurenti. Valle: Rio Calima. Calamita palmatus. 32 (type locality not known). Shaw.— Liu. 1897. p. p. 1960. p. 174. 1931.— Andersson. Boulenger. CJG 2258. 163. 152313-5. p. and webbing dusky between fingers and toes. 1799. 17. Hyla maxima. 1933. p. p.— Wagler. p.— Gorham. 26 (type locality.— 300. 1963. . 75. p. EPN 3080. p. p. Gorham. 173. Rio Cayapas. Tschudi. EPN 4280. 1858. 66. 1873a. Andersson. p. 1803.— Cope. Venezuela). 217. OV 113. 1931. Hyla miranda-ribeiri Melin. Hyla boans. 97. 99. p. 1963. 200. Rana — Schneider. 2983-4. p. 207. 2913-5. 3040. Cayenne) 1803. 34. Nieden. — — 1941. p. 1820. 1958. p. p.147-51 (types of Hyla 2900-11. 1758. p.— Goin. p. p. 1055. p. 3038. 137. 1919. upper Rio Sinii. 213 (type locality. Rivero. 17. 79.— Rivero. 1896. p. 1958. and the webbing between fingers and toes dusky to dark gray. San Lorenzo. 1904.— Gorham. 7418. p. 1964. 118. 1882a. 23. Nanegal. 1830. p. 20 (type locality. — Gmelin. 22. Hyla lactea Lonnberg.— Lutz and Kloss. 22. hands and feet fully webbed.— Beebe. Hypsiboas palmata. p. 1799. Rio Raposo. 1830. Ecuador. p. Carolina and Virginia). 1802. rosenbergi) . p. 1952. p. 1841. 1923.— Thornton. Cachabi. 349. Brazil). above Gamboa). 1768. PANAMA: Alhajuela (10 mi. 1961. 544. 3034-6. Pacto. 1900. — 36.— Gunther. 218. p. 89. 1961. p. 509. Rfo Vaupes.— Miranda-Ribeiro. Peters. 96. 650. USNM USNM 151385-7. ECUADOR: Esmeraldas Province. p. 1900. p. 1867. p. 2 (type locality.— Werner. Amazonas. — 1820. . p. 13 (substitute for Hyla boans L. 17. 72. 1945.— Noble. Pichincha Province. northwestern 10090. triangular dermal appendages on heels.). 33. 188. 109. 1933. H. 1931. 2936-44. Calamita maxima. 2460-2. 1963. p.— FROGS OF COLOMBIA — COCHRAN AND GOIN 199 Specimens Examined COLOMBIA CNHM C6rdoba: SocarrS. Merrem. — 1858. p. Baja Rio Toachi.— Crawford and Jones. EPN 4027. 61167. 1935.

loreal region concave and very oblique. a low indistinct glandular ridge encircling upper part of tympanum. NATIONAL MUSEUM BULLETIN — CNHM 2 88 54789.8 mm. Body elongate. Colombia. \ series.. heel reaches beyond tip of snout. heels overlap slightly. third and fifth toes subequal. slightly projecting. Eye moderate. dirty tannish gray below. a very indistinct tarsal ridge. Tympanum very distinct. with most of them clustered in middle of the back dorsal pattern consists of a narrow tip of . Can thus rostralis well defined. their distance from end of snout about one-fourth that from eye. no vocal sac. Toes completely webbed. Head and body. are scattered over the back. when hind leg is adpressed.. broadly triangular when viewed from above. 30. the upper jaw extending a little beyond lower. heel-to-toe. that of belly and lower surface of thigh uniformly granular. skin of throat and chest smooth. the upper lip flaring out strongly below it. separated from each other by an interval equal to about one-half their distance from eye. when limbs are laid along the side. The median stripe that extends from snout to about the sacral hump.S. the web on fourth toe reaching the base of the disk. a distinct oval inner but no apparent rounded outer metatarsal tubercle. Some dozen small brownish spots. tongue three-fourths as wide as mouth. about one-half the diameter of eye.. a short broad triangular dermal appendage on heel. Skin of head not co-ossified with skull. which is relatively wide and slightly greater than distance between nostrils. mm. nostrils more lateral than superior. No patagium. when hind legs are bent at right angles to body. 32. —Pale brown above. its posterior border very slightly free and very shallowly notched. adult female.6 mm. broadly cordiform. femur. oblong choanae. the web on third finger reaching base of disk on lateral side and middle of penultimate phalanx on medial side. fairly prominent. head length. lying close together between Vomerine teeth in two heavy / the large. a few millimeters in diameter. interorbital distance half again as great as width of upper eyelid. Snout large.3 Color in alcohol. — Dimensions. rounded in profile. from Pueblorrico. roof of skull not exostosed. 90 mm.200 U.3 mm. a low ulnar ridge. reaching to middle of disk of third which more than covers the tympanic area. knee and elbow overlap. opening. its diameter not quite as great as its distance from nostril. head width. palpebral membrane reticulate. tibia. fourth finger longer than second. 27. Skin of upper parts smooth. Description. 47. hand. no traces of a skinfold across chest.2 mm.. separated from eye by a distance nearly equal to one-half its own diameter. rudiment of a pollex not pronounced.. in postaxillary region but little narrower than greatest width of head. disk of fourth toe slightly larger than the tympanic area. 63.. where it fades out. 49. Fingers webbed.1 mm. Caldas.

however. preoccupied boans Latreille. In the meantime Laurenti (1768. 1923. at most. Crawford. Boulenger. The series at hand. 1931. shows considerable variation in pattern. Peters. 32) gave the name maxima. although it does have an in Venezuela abortive dorsal pattern. p. In addition to size. and webbing of hands and feet are a very dark gray that approaches black.7 to 107. compared to five adult specimens (taken at random from the series before us) that range from 87. — Remarks. and forearms. dusky throats) and their fingers and toes are much more slender and delicate. 17). which became adopted as the name for the species and was used widely for many years (Giinther. from Leticia. p. Rather indistinct crossbars occur on the dorsal surfaces of thighs.FROGS OF COLOMBIA —COCHRAN AND GOIN 201 above shoulder region. Miranda-Ribeiro. The largest specimen at hand. Geographically speaking. particularly those from the Rio Atrato. 1931. narrow. underside of feet. Medem on the Serrania de La Macarena. but the triangular dermal heel appendage and the reticulate palpebral membrane seem to indicate that such specimens definitely are maxima rather than is — rosenbergi. 1945). p. 1858. There can be no doubt that this frog was first named Rana boans by Linnaeus in the tenth edition of his Systema Natura. We have examined the specimen from San Fernando de Atabapo (USNM 80652) mentioned by Rivero (1961. the specimens that show the most divergence from "typical" maxima are two frogs collected by Dr. Along each side between the axilla and groin a series of vertical dark bars. measures 115 millimeters in head-and-body length. until 1900 when it was resurrected by Andersson (1900. these becoming more distinct posteriorly in region of the groin. p. Variation. That name was overlooked.2 millimeters in length. feet. with the webbing on the fingers being more reduced and the webbing itself being much darker than it is in the average maxima. shanks. Most of the specimens are brownish with a dark. median dorsal stripe running from the tip of snout to above the urostyle. On 1801. . 1882. 1872. Nieden. while structurally quite uniform.7 and 73. Cochran (1955. Andersson. 80) used the name albopunctata for the frog of southern Brazil allied to lancijormis of Colombia which had for so long been known as boans. Posterior surfaces of thighs. the basis that boans Linnaeus. A few. 1926. these two specimens differ in that their throats are quite dark (whereas most examples of maxima have light throats.3 millimeters in head-and-body length. Noble. seem to have the webs paler than others. 1758. 97) and find that it is structurally typical maxima. These specimens are 75. or.

maxima. Ill) argued that. is 1758. dermal appendages on the heels. Art. p. 1758. as unavailable (Code. So far as we know. 1955. p. 1768. 23. the only recent major work to use the name boans for what we here call maxima is that of Rivero (1961. and These frogs generally are tannish to brownish in dorsal ground color and often have a narrow dark median stripe down the back. London. faber. p. and on this basis we consider boans Linnaeus. This precise wording now available (International Code of Zoological Nomenclature. Although these frogs are all closely related they are not just races of a single species. called albopunctata With the assignment there are still of wavrini and miranda-ribeiri to synonymy three closely related. becomes available for the frog that name Cochran from southern Brazil (Cochran. 25). for two of them. and it also occurs in the Rio Atrato drainage of Choc6 and has penetrated southward to the valley of the Rio Dagua in northern Valle. 2) quite properly diagnosed it as separate from jaber and rosenbergi but apparently he did not compare it with maxima.b). large. The names of two recently described forms seem to be synonyms of H.b. a nomen oblitum pending the precise wording in the revised Code. 23. 1758. maxima is wide-ranging through the Amazon Valley from French Guiana to the eastern slope of the Andes. a specimen of maxima without any doubt. Art. the specimen in the British Museum proved to be.S. Parker (1936. in our opinion. p. for the frog so long With the suppression of the name ii) and use the name maxima that name. Latreille. 80). all characteristic of maxima. 1953. While rosenbergi is a frog of the Pacific coast in South America (one record from C6rdoba. and that the absence of records for coast of Colombia is due simply much to lack of collecting. On direct comparison with specimens of rosenbergi. Colombia). iaber. 1961. on the basis of the Copenhagen decisions (Hemming. and maxima. p. it would be better to leave the name maxima for the frog so long known under that name and to consider the boans Linnaeus. namely. 2 88 Goin and Layne (1958.202 NATIONAL MUSEUM BULLETIN U. rosenbergi. have overlapping ranges in the Rio Dagua drainage in southwestern Colombia. We suspect that maxima most likely reached the Rfo Atrato drainage by way of the Atlantic coast of Colombia rather than across frogs that are apt to be confused. and dusky webbing on the hands and feet. 96). the boans Latreille. Examination of the specimen that Melin used as the type of Hyla miranda-ribeiri demonstrated that it likewise is a maxima. very similar appearing tree maxima. the Andes. maxima and rosenbergi. It has the reticulate palpebral membrane. We have examined the paratype of Hyla wavrini and it seems to be maxima without any question. 1801. known under boans Linnaeus. of the northern .

73797. BRAZIL: Rio Vaupes. p. BM 1936. Vaupes: Rio Cuduyari. Payamino. GUIANA: Cayenne. 152630-1. MLS 49. geographica geographica fingers with just a trace of web H. 99— Miillcr. Rio Llushin. who call it "cagua. JAP 3746. Rio Villano. EPN 1753. ANSP 25310.— Gunther. 3745. p. fingers . 1658-61. Boca de Guataracu. The two Colombian species may be distinguished by the following key: heels a . 152131. Palpebral membrane reticulate. 151998-9. MLS "b. 152101-4. EPN 1548-52. EPN 1829. Moderate-sized tree frogs with elongate dermal appendages on the and reduced webbing between the fingers. ex. Cabeceras del Rio Bobonaza. Puerto Narifio. Rio Raposo. fig. Randal 152132-3. 152189. 303.67. Guacaya. Cabeceras del Rio Capahuari. 7896. near El Refugio.29. Palpebral membrane clear.2. Serranfa de Baud6. JAP 2233-4. Cafio 8534 (2). H. 99. p. FROGS OF COLOMBIA — COCHRAN AND COIN 203 Dr.— Lutz and Kloss. EPN 1647-8. EPN 1357. 151385-6. "Region de Puyo. 1932. 152319. Caldas: Pueblorrico— Santa Cecilia. upper Orinoco. 30575. C6rdoba: Upper Rio Ure. Morelia. Leticia. 1964a. p. Hyla geographica Spix.23. EPN 1615. Taracua. Montalvo. Rio Popoyaca. ANSP 25314. Rio Pucayacu. Meta: Near mouth of Caiio Losada. EPN 1345. Rio Apaporis. Valle: BM 1908. 152134-5. Macanal. CJG 2464 (6).12. UF 8532. 28051. 1 (type locality unknown). 1 a2 about one-half webbed. Cabeceras del Rio Solis. 1858. EPN 1446.— Rivero." 15220. 1856. Hyla geographica geographica Spix Plate 27a-c 1824. GEOGRAPHICA group 4. 78143-4. OrceVV. .5.10. forest between Macarena and Guayabero. Serram'a de La Macarena. — Burmeister. 71216. 1913. Loreto. calcarata . 151989. Villa Arteaga. Jirijirimo.1 [1947. 466 (type of Hyla miranda- USNM USNM USNM NM ribeiri) ECUADOR: Napo-Pastaza Province. EPN 1715-6. 2023-4. 54785-91. MLS 194. Alto Napo. EPN 3557. EPN 4044-5. 1922." Boyaca: Guaiearamo. Amazonas: 8533 (2). EPN 2451-2. MLS 53. south of Buenaventura.30] (paratype of Hyla wavrini). 152195.116-7. 170. p. Antioquia: Urabd. 146263. 152194. USNM 80652. Rio Conambo. Rio Orteguaza. Choc6: Condoto. "El Mico. Quibdo. MHNP FRENCH VENEZUELA: San Fernando de Atabapo. of the Escuela Politecnica Nacional tells us that is eaten regularly by the Indians. . 1952." EPN 1678. p. 22.10.— Gorham. 649.. Caqueta: Florcncia." maxima Specimens Examined COLOMBIA USNM 152130. 1963. 4618 (type of Hyla palmala). 152128-9. Chicherota.11. Sautata. JAP 3732. Villavicencio. 1637-8. MLS 15. 63894-6. EPN 1722. La Providencia. USNM USNM USNM MCZ USNM CNHM MZUM CNHM USNM CNHM BM CNHM USNM USNM USNM USNM USNM CNHM MCZ Los Mangos. 11622-3.

Cochran. p. which is only slightly greater than distance between nostrils. 5.— Miranda-Ribeiro. Fingers more than one-half webbed. 1945. an adult female from Guaicaramo. This subspecies may be distinguished from all Colombian Hyla except H. 1926. Andersson. p. appendiculata Boulenger. inference). — Hyla punctatissima appendiculata. tongue three-fourths as wide as mouth opening. by 1927. 349 (type locality. and the fingers about one-half webbed. 180. It may be readily distinguished from H. 600. about three-fifths the diameter of eye. U. separated from each other by an interval equal to about two-thirds their distance from eye. Description. 1901. Canthus rostralis well defined. no projecting rudiment of a pollex. a distinct ulnar ridge. 64. Parker. broadly triangular when viewed from above. p.— Canelos.— Nieden. separated from eye by a distance about equal to one-third its own diameter. Snout moderate. p. Brazil. 1955. 220. considerably projecting. interorbital distance half again as great as width of upper eyelid. Colombia. 1933. Parker. — . MLS 64. 1935. Ecuador). its diameter equal to its distance from nostril. web on fourth toe reaches base of penultimate phalanx. 195 (type locality. nostrils more lateral than superior. p. A moderate-sized tree frog with the palpebral membranes reticulate. — — p. just reaching to middle of disk of third which covers about two-thirds the tympanic area. Toes three-fourths webbed. third and fifth toes subequal. p.S. p. — Diagnosis. a spur-like dermal appendage on the heel. 99. Vomerine teeth in two heavy angulate series which touch on the midline and lie between the large. Eye not large. 1882. the upper lip flaring out rather strongly below it. oblong choanae. Tympanum very distinct. p. palpebral membrane reticulate. 511. p. maxima by the reticulations on the palpebral membranes.— Baumann. p. here restricted.— Werner. 70. nor particularly prominent. broadly cordiform. 93. NATIONAL MUSEUM BULLETIN 288 Hyla — Lutz. fourth finger considerably longer than second. its posterior border fused and very shallowly notched. Hyla spectrum Reinhardt and Liitken. the upper jaw extending slightly beyond lower. p. 1923. somewhat truncate in profile. maxima by the much larger appendage on the heel and by having the fingers only one-half webbed. 1933. loreal region concave and oblique. 1897. disk of fourth calcarata. 1912. Hyla geographica geographica. 1882a. on all other toes it reaches to base of disk. the web reaching base of the penultimate phalanx of the fourth finger on the lateral side and extending thence as a narrow margin on the medial side. 301. their distance from end of snout about one-third that from eye.204 1862. In the structure of the dermal heel appendage it is most like Hyla but it differs from that species in having the palpebral membranes reticulate and in having the fingers about one-half webbed rather than with just a trace of web. Boyaca.

while in MLS 64 the tympanum is little more than half the diameter of the eye and is separated from it by one-half its own diameter.7. The entire side. Dimensions. head length. in postaxillary region a little narrower than greatest width of head. On the very dark dorsum an even darker blotch can be distinguished. heels overlap greatly. head width. the dorsal markings are broad rather than pencil-like. A dark frog. 27.7 mm. lightcentered vertical bars. a distinct oval inner but no distinct outer metatarsal tubercle. The ventral surfaces are immaculate. when hind leg is adpressed.13. no trace of a skinfold across chest. 53. in the MLS specimen they meet on the midline. knee and elbow overlap considerably.. We strongly sus- — — — . roof of skull not exostosed. femur. Body somewhat elongate. skin of throat and chest smooth. In the cotype of appendiculata.7 mm.2 mm. pencil-like. quite distinct glandular ridge encircling upper part of tympanum. then passing downward to angle of jaw. is marked with distinct. that of belly and lower surface of thigh uniformly and finely granular. 15.2. The patterns of the sides and the thighs seem to be about the same in both specimens. from axilla to groin. which is quite dark dorsally.. when limbs are laid along the side.4 mm. a narrow..FROGS OF COLOMBIA —COCHRAN AND GOIN 205 toe covering about one-half the tympanic area. When the above-described specimen (MLS 64) is compared with one of the cotypes of appendiculata (BM 74.7]) from the interior of Brazil.7 [1947.5 mm. 38 mm. Color in alcohol. hand. transverse markings across the tan dorsal background. an elongate dermal appendage on heel. Similar bars are present on the anterior face of each thigh. No patagium. heel-to-toe. 19. no vocal sac. adult female.. The cotype is somewhat faded and at present the dorsal pattern consists of narrow. in the MLS specimen.. Less-distinct narrow cross-bars are also present on the shanks. Skin of upper parts very finely granular. These continue over the dorsal and posterior surfaces of the thighs but are less conspicuous there due to the more intense pigmentation of the ground color. from the middle of which two posteriorly projecting horns diverge. the two series being continuous. Head and body. The rami are in turn connected by a narrow bar at the level of the sacral hump.16. This blotch anteriorly has a narrow ramus that extends to the hind corner of eye on each side and thus makes a Y-shaped pattern. 29. Posteriorly the blotch again divides to send a ramus to each lateral extension of the sacral hump. Skin of head not co-ossified with skull. Variation. when hind legs are bent at right angles to body. the tympanum is about three-fourths the diameter of the eye and is in contact with it. In the cotype there is a definite gap between the vomerine series. 20 mm. several differences can be noted.. tibia. a weak but distinct tarsal ridge. heel reaches almost to nostril.

ger. . Bobo2733. Lutz and Kloss. Most of the adult specimens from the northern part of the range within the range of variation discussed above.10-1. MLS 64 (5). Lutz and Kloss (1952. naza. 354.166-8 (cotype3 of appendiculata). 200. ANSP 24401. There is a beautiful bright yellow spot on each heel." EPN 1689. BM 69. 306. Cope. p. Para. NATIONAL MUSEUM BULLETIN 2 88 pect that the differences in the tympanum and (possibly) in the teeth are due either to sex or to geography. p. EPN 1349-50. 1862. 206 U. 1904. p. p. p. Hyla calcaratus—Cope. "afluente del Rio Pastaza. 1963. 94. p. 1904.5." EPN (two unnumbered specimens) Napo. de 4115-7.— Beebe. Alto Oglan. Remarks.12. 208. Rio Villano. Santarem. USNM 152100. 1952. and the belly is uniformly. p. 647. interior of Brazil.22. 34." 1333. 20. EPN 3519. Rio Conambo. 660 (type locality. 359. EPN Hyla calcarata Troschel Plate 27d-f 1848. 1931. Hyla fasciata— Lidth.13. BM 80. EPN 1521. BM 74.16. Chichirota.11. Boulenger included a specimen from Bahia (BM 69. 1919. ECUADOR: Napo-Pastaza: Canelos. 3758.5. JAP EPN EPN EPN EPN EPN 1356. Cundinamarca: Eastern base of cordillera of Bogota.— Gorham. 1862b. 1867. 1352-3.25 That specimen [1947. Rio Rutuno.7. — Boulen— 1945. p. 1867. however. Baja Bobonaza. BM 75. . 5). as it is much paler than the northern form of H.— Nieden. but not thickly. Loreto. p. Boca del Coca.12. Alto Rio Pucayacu. Sumaeo. p. British Guiana). p. geographica dorsally. p. Their young specimens were the slate-blue color of early metamorphosed forms.10. Hyla calcarata Troschel. 2501. alto Pastaza. Rio Pindo. Alto Rio Pucayacu. "N.11. Rio Llushin. 1882a.. (unnumbered specimen).25. Parker (1933. "region de Puyo.3. EPN 3889.S. 69 — Hypsiboas fasciatus. Rio Verde. Copataza. but since that species lacks the reticulations on the palpebral membrane it should be readily separable fall from young H. but always near still or slowly flowing water. Sarayacu. speckled.2.6]) in the series of cotypes of appendiculata. p. Specimens Examined COLOMBIA Boyaca: Guaicaramo. probably should be referred to II. EPN 1486-7. geographica punctatissima. He describes young individuals as having a pattern similar to that of adult calcarata. 2036.3.234 (cotype of appendiculata) San Jose Viejo de. reports that the young have quite an ontogenetic change in color and pattern. 649) point out that all their specimens from the upper Amazon region were taken in the daytime on the forest floor. 1923. BM 80. BRAZIL: All cotypes of appendiculata: Bahia. but even at that early age they had the reticulations on — the palpebral membranes which are characteristic of the species. JAP 3797.— Crawford. Arapicos.— Andersson.7. geographica geographica.

— CNHM appendage on heel. oblong choanae. not projecting. separated from eye by a distance about equal to one-third its own diameter. Snout large. Body somewhat elongate. Serrania de La Macarena. a very distinct. tongue two-thirds as wide as mouth opening. then distance from end of snout about one-third that from eye. an adult female from Rio Guapaya. just reaching to disk of third which covers about two-thirds the tympanic area. it can be distinguished by the absence of palpebral reticulations. broadly cordiform. the upper lip flaring out below it. and without reticulations on the palpebral membranes. Meta. From H. heel reaches beyond tip of snout. interorbital distance slightly greater than width of upper eyelid. which it closely resembles. no projecting rudiment of a pollex. 81333. lying close together between the large. palpebral membrane not reticulate. maxima. Superficially.FROGS OF COLOMBIA Diagnosis. loreal region concave and oblique. —A — COCHRAN AND GOIN 207 moderate-sized Hyla apparently allied to H. a rather narrow glandular ridge encircling upper part of tympanum. heels overlap greatly. but it can be distinguished from that species by the greatly reduced webs between the fingers. its posterior border fused and with but a trace of a notch. elongate dermal graphica. Tympanum very distinct. H. nostrils more lateral than superior. No patagium. the upper jaw extending slightly beyond lower. a distinct oval inner but no distinct rounded outer metatarsal tubercle. third and fifth toes subequal. when limbs are laid along the side. rounded in profile. Vomerine teeth in two heavy series. with dark markings in region of groin posterior faces of thighs. partialis. fourth finger considerably longer than second. somewhat pointed when viewed from above. rosenbergi it differs in having dark markings in the groin and on the thighs. knee and elbow considerably overlap. geographica and young H. when hind leg is adpressed. geo- and on anterior and with exceedingly well-developed dermal spurs on heels. Colombia. Description. Toes slightly more than one-third webbed. about one-half the diameter of eye. its diameter equal to its distance from nostril. Canthus rostralis slightly defined. Fingers with a trace of web at base. . disk of fourth toe covering about one-third the tympanic area. and a well-developed dermal spur. Skin of upper parts smooth. calcarata closely resembles H. when hind legs are bent at right angles to body. the web on fourth toe not quite reaching the base of the penultimate phalanx. not particularly prominent. in postaxillary region a little narrower than greatest width of head. skin of throat and chest smooth. Eye moderate. which is in turn slightly greater than distance between nostrils. a distinct but not conspicuous ulnar ridge. From young H. a smaller tympanum. separated from each other by an interval equal to about two-thirds their distance from eye. tarsal ridge small but distinct.

. A study of the type of fasciata (BM 58.6 mm. 208 U. tibia. CNHM 69715. p. 152317. roof of skull not exostosed.1. head width.6 mm. hand. New River. 17.1. head length.8 mm. 34) have synonymized Hyla fasciata Giinther and Hyla calcarata Troschel. Light brown above. 306. Remarks. Lutz and Kloss (1952. tympanum too significant to permit synonymizing the two forms.2 mm.. BM . 359. 1931. anterior and posterior and inner face of shank heavily marbled with black. p. heel-to-toe. BRITISH GUIANA: 1939. USNM 2353. while fasciata is a more montane form occurring in the Andes proper. The posterior part of side.22) shows that it differs from the specimens of calcarata at faces of thigh... Specimens Examined COLOMBIA Caqueta: Rio Mecaya. CNHM 81333-4.4. femur. While we have not examined very much material. 25.61. Whether the two are simply geographic races of a single species cannot be determined until additional material has been studied. 28. Skin of head not with skull. loreal only one-third rather than onehalf the diameter of eye. p.2.. While there can be no doubt that these are closely related forms. — hand more strongly defined. This locality is just across the river from Leticia.7.4 mm. It seems to us that these differences are region as follows: the canthus rostralis much less oblique. CJG Putumayo: Puerto Asfs on upper Rio Putuuiayo. 14.25. USN 152279. yellowish gray below. digital disk of third finger covers entire tympanum instead of only two-thirds of it. Several workers (Boulenger. 48. BM 1936. Meta: Serranfa de La Macarena. A narrow dark stripe lies on each canthus. we do not believe they may be appropriately synonymized. NATIONAL MUSEUM BULLETIN 288 that of belly and lower surface of thigh uniformly and finely granular no vocal sac. Head and body. Color in alcohol. p. Immaculate below. dermal appendage on heel short and triangular instead of long and spur-like. Dimensions. 647) report that their specimen from Benjamin Constant. from specimens seen it appears likely that calcarata is a lowland form ranging from the Guianas to the eastern slopes of the Andes. Nieden. we have Colombia. Crawford. and a similar stripe originates at tip of snout and passes posteriorly between the eyes and fades out no trace of a skinfold across chest. web on fourth toe extends to distal third of penultimate phalanx instead of failing to reach the penultimate phalanx. 1923.S.35.3 mm.5 mm. Vichada: Amanaven. Villavicencio.. 17.. 1882. co-ossified — — above the tympanum. 35. Matanuki River. Brazil. was found between two bromeliad leaves on a fallen tree.

lying close together on a level with the posterior borders of the large. 309 (type locality. nostrils lateral. — COCHRAN USNM 118709. Snout large. third. large — beyond lower. separated from each other by an interval equal to about two-thirds their distance from eye. Canthus rostralis strongly defined. near Antioquia-Bolivar border. considerably projecting. interorbital distance slightly greater than width of upper eye lid. 5. Colombia. diameter. Gorham. separated from eye by a distance nearly equal to one-half its to disks of second. its diameter equal to its distance from nostril. 22. Abitagua. its posterior border fused and with just a hint of a notch. hands and feet. Rio Cauca. . p. — p. loreal region concave and somewhat oblique. and transverse patches of vomerine teeth of this species serve to distinguish it easily from any other Colombian Hyla. Puyo. Eye large. tongue two-thirds as wide as mouth opening. extensively webbed hands and feet. and sacral diapophyses not appreciably expanded. very large. Diagnosis. Description. MLS 267 (holotype) from Rio Cauca. 516. Fingers webbed to base of penultimate phalanx own but half a disk longer than second and reaching middle of disk of third which much exceeds the tympanic area and nearly covers the eye.FROGS OF COLOMBIA ECUADOR: Oriente. immensa of Costa Rica is the only other species we know of that is a member of this group. the upper lip flaring out distinctly slightly their distance below it. broadly rounded. third and fifth subequal. fourth finger to . PHANTASMAGORIA group Very large tree frogs with vomerine teeth in transverse patches. which is relatively wide and slightly greater than distance between nostrils. ulnar ridge well developed with scalloped margin. from end of snout one-third that from eye. Vomerine teeth in two long. broadly U-shaped when viewed from above. Toes completely webbed to all disks. — An immense Hyla are extensively with the The with enlarged hands and feet that the cranial derm not co-ossified webbed and with skull. At present. and fourth. heavy transverse series. Tympanum very distinct. disk of first. about one-half the diameter of eye. H. JAP 1956. AND GOIN Napo 209 Pastaza. 1963. Hyla phantasmagoria Dunn. rounded choanae and extending laterally to terminate behind the median third of the choanae. the upper jaw extending large size. near border of Antioquia and Bolivar) 1944c. palpebral membrane not reticulate. Hyla phantasmagoria Dunn Plate 28a-c 1943. very prominent. a pronounced projecting pollex covered with soft flesh. p. truncate in profile.

femur. chest. 60.8 mm. Skin of head not co-ossified with skull. Head and body. no dermal appendage on heel. 41. wellof fourth toe developed tarsal ridge.8 mm. laid No along the legs are with prominent. Elsewhere the dark pigment is so concentrated that it leaves no discernible pattern. Dimensions. The only possible conclusion was that the two species are very closely related. when limbs are considerably. a strongly developed glandular ridge passing above upper part of tympanum and continuing posteriorly to above arm.. Duellman brought the type of Hyla immensa Taylor from Costa Rica to Gainesville. very dark brown with no discernible pattern although slightly paler above sacral region. Florida. heels patagium. head length. when hind overlap greatly. tibia.S. no vocal sac. while on the undersurfaces of thighs it appears as larger blotches that are separated by narrow bands of lighter area. roof of skull not exostosed. side. including the arms and legs with a well-developed patch on each heel.3 mm. adult female. in postaxillary region a little narrower than greatest width of head. skin of throat. H. and lower surface of thigh uniformly and coarsely granular. a distinct oval inner but no distinct rounded outer metatarsal tubercle.. 50... 84. somewhat paler than above but still quite dark. wart-like tubercles. heel reaches almost to nostril. Body not elongate.5 mm. and he and the junior author compared it with the type of H. 42 mm. these tubercles conspicuous on all dorsal surfaces. —We do not understand how Dunn came to describe having the skin of the head co-ossified with the skull. phantasmagoria.7 mm. The general effect is that of a dark. carri group Large tree frogs with reduced webbing between the fingers and a dorsal pattern of dark blotches bordered with light margins. but we do not wish to make this assignment after examining only two specimens. Below. 6. On chin and throat the dark brown pigment is in a more or less vermiculated pattern. daresignata of Brazil apparently belongs in this group. 35. chocolately brown frog with slightly lighter — — ventral surfaces. head width. . 109. knee and elbow overlap bent at right angles to body.3 mm. elongate. when hind leg is adpressed. hand. Skin of upper parts highly glandular. belly.210 U. heel-to-toe. this species as the skin is fused to it. Above.. for we have made a slit in the skull cap and can move a needle completely over the top of the skull without finding any evidence that Remarks.. It is very possible that they represent different races of a single species. NATIONAL MUSEUM BULLETIN 2 88 somewhat larger than the tympanic area. no traces of a skinfold across chest. Color in alcohol.

Snout moderate — Figure 19. . in Huila. Hyla carri. new AND GOIN 211 species Figure 19 — CNHM — Holotype. somewhat squarish choanae. nearly round. collected at Acevedo. from which it can be differentiated by its more heavily spotted dorsum.— FROGS OF COLOMBIA Hyla — COCHRAN carri. along the Rio Suaza. The species is not similar to any other species known in Colombia. larger tympanum. Colombia. Diagnosis. 69702. Description of holotype.thirds as wide as mouth opening. CNHM 69702 (same size). reduced webs between the fingers. its posterior border fused and very shallowly notched.400 meters. and a pattern of dorsal dark blotches bordered by light margins. 1. lying close together between the moderate-sized. Vomerine teeth in two -^ ^> series. and lack of dark anal spot. It is perhaps most closely related to Hyla claresignata of Brazil. A large Hyla with the vomerine teeth in two -^ ^ shaped patches between the somewhat squarish choanae. tongue two. holotype.

8 — mm. which is in turn about equal the distance between nostrils. truncate upper jaw not extending appreciably beyond lower. No patagium. Skin of head not of with skull. 22 mm. when hind leg is adpressed. The chin. NATIONAL MUSEUM BULLETIN 288 somewhat rounded when viewed from above. then continuing as narrow margin to base of disk. Toes nearly completely webbed. and chest are fuscous. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. in postaxillary region a little narrower than greatest width head. but the blotches on anterior portion of trunk and on top of head are smaller than the more posterior ones and are not margined with a lighter tone. its diameter equal to its distance from nostril. 23. throat. when hind legs are bent at right angles to body. reaching to middle of disk of third which just covers the tympanic area.3 mm. Canthus rostralis well defined.. separated from eye by a distance equal to one-half its own diameter. no tarsal ridge. when limbs are laid along the side. about two-thirds the diameter of eye. in profile. Head and body. a dirty gray. no projecting rudiment of a pollex. co-ossified 50. heels overlap greatly. their distance from end of snout about one-third that from eye. considerably projecting. palpebral than width of upper eyelid. tibia. interorbital distance slightly greater middle of antepenultimate phalanx. a strongly developed narrow glandular ridge encircling upper part of tympanum. and feet the dorsal blotches are outlined with a narrow pale margin of tan. 38. fourth finger considerably longer than second. third and fifth toes subequal. femur. the web on fourth toe reaching the middle of the penultimate phalanx. 64. A moderate brown frog with a dorsal pattern consisting of dark brown blotches. Fingers webbed at base. no dermal appendage on heel. disk of fourth toe covering about four-fifths the tympanic area. no traces of a skinfold across chest..212 U-S. and here there are a number of smaller. skin of throat and chest very finely granular.3 mm. web on third finger reaching membrane not reticulate. the upper lip flaring out moderately below it.— . that of belly and lower surface of thigh uniformly and rather coarsely granular. no ulnar ridge.. knee and elbow overlap considerably. black spots. head width. the nostrils more than superior. Eye moderate. rather distinct.3 mm. Color in alcohol... Along each side between axilla and groin the ground color is much lighter than elsewhere. On posterior portion of trunk and on forearms. hand. Body moderately elongate. head length. Dimensions. 22. no vocal sac.. loreal region concave and somewhat oblique.5 mm. roof of skull not exostosed. Tympanum very distinct. heel-to-toe. 33. separated from each other by an interval nearly equal their distance from eye. heel reaches beyond tip of snout. adult female. not particularly lateral prominent. legs. Skin of upper parts smooth.1 mm.

1882a. separated from each other by an interval equal to about two. rounded when viewed from above. a gravid female packed with pigmented we have seen of this species. and we know no specimen from the Andes that resembles it. reduced webbing between the fingers. Canelos). throat and chest dusky but not distinctly spotted. species in this group of the following may be distinguished key Faded mottled pattern on back. 1. 362 (type locality. — Diagnosis. the upper lip flaring out below it. loreal region concave and rather oblique.- —COCHRAN AND GOIN 213 holotype. Ecuador. buckleyi distinguished is most closely allied to cabrerai. 1923. buckleyi group Moderate-sized Hyla with well-developed patagia. 267. and choanae moderate in size and somewhat oblong rather than small and rounded. —CNHM Nariflo. thirds as wide as 337-262—70 15 . fingers about one-half webbed H. Eye moderate. here restricted. lying close together between the moderate-sized.thirds their distance from eye. throat and chest regions dusky to distinctly spotted. broadly rounded. Vomerine teeth in two short AA series. Colombia. Description. throat and chest dusky but not distinctly spotted. Can thus rostralis well defined. buckleyi Distinctly marked above. not particularly prominent.third that from eye. and reduced webbing between the fingers.: FROGS OF COLOMBIA —The Remarks. Snout moderate. fingers webbed only at base H. palpebral membrane not reticulate. 1961. — Nieden. A moderate-sized Hyla with fairly well-developed patagium. their distance from end of snout about one. is the only individual of given us. a3 . Osteocephalus buckleyi. Hyla buckleyi Boulenger. its posterior border very slightly free and unnotched. oblong choanae. 13. its diameter equal to its distance from nostril. from Rio Rumiyaco. tongue two- mouth opening.000 meters. from which by the reduced webbing between the it fingers may be and the lack of distinct spots on throat and chest. throat and chest heavily sprinkled with small distinct spots. p. p. We dedicate this new species to our friend Archie Carr in recognition of his contributions to tropical biology and for the help he has eggs. cabrerai Hyla buckleyi Boulenger Plate 28d-f 1882.— Goin. the upper jaw extending but little beyond lower. 7. truncate in profile. nostrils more lateral than superior. The two Colombian with the aid a1 . p. considerably projecting. H. 54765. faded mottled pattern on the back.

tympanum.5 mm. no vocal sacs. 48. when hind legs are bent at right angles to body. Our notes made at time of is com- parison are as follows: CNHM Most of the differences can be laid to differences in preservation. no projecting rudiment of a pollex.4 mm. Toes slightly more than one-half webbed.2 mm. Tympanum very distinct. if it is studied caresmooth. The entire undersurface is a dark cadaverish gray. reaching to middle of disk of third which covers about one. Remarks. Dimensions. The CNHM .S. femur. The specimen has the skin harder and drier while the cotypes have the skin soft and specimen is much darker. no dermal appendage on heel but three definite warts arranged in a triangular position on each heel. heels overlap considerably. and body. roof of skull very lightly exostosed. Each loreal region has a mottled area. a . Each thigh and shank is crossed by three rather wide crossbands of dirty brown with less-distinct bands on the feet.. somewhat transverse smudge above the sacral hump..4 mm.. knee and elbow overlap. which wide and greater than distance between nostrils. 17. and hands. the web on fourth toe reaching the base of the penultimate phalanx.. disk of fourth toe covering about one-third the tym- panic area.. separated from eye by a distance equal to one-fourth its own diameter. —The specimen from Nariiio (CNHM 54765) was directly compared with the series of cotypes in the British Museum and found to be conspecific with them. arms. Skin of head not co-ossified with skull. hand. is relatively Fingers slightly webbed at base. about three-fourths the diameter of eye. that of belly and lower surface of thigh uniformly granular. head width. There is a narrow. interorbital distance slightly less than width of upper eyelid. but even so. heel reaches just beyond tip of snout. fourth finger considerably longer than second. mm. A series of small blotches are scattered along each side between the axilla and groin. 34. no tarsal ridge but several small warts along the posterior edge of foot.third the tympanic area. third and fifth toes subequal. —A dirty brown frog with darker mottled brown blotches on back and side. 14 Color in alcohol. adult female. Skin of upper parts very finely glandular. in postaxillary region a little narrower than greatest width of head when hind leg is adpressed.4 —Head 17. heel-to-toe. distinct glandular ridge encircling upper part of of throat and chest fairly skin smooth. no traces of a skinfold across chest. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Body moderately elongate. no ulnar ridge.214 NATIONAL MUSEUM BULLETIN 288 U. 27. tibia. when limbs are laid along the side. A patagium extends from the back of upper arm to side of the body. There is a large smutty blotch on the back that extends from between the eyes nearly back to the sacral hump.7 mm. head length.. mm. 24.

the upper jaw extending but slightly beyond lower. NariSo: Rio Rumiyaco. which is relatively wide and slightly greater than distance between nostrils. by I. rounded when viewed from above. Sarayacu. October 17. the upper lip flaring out below it. 1951 by I. Brazil. Diagnosis. lying between the moderate-sized. . new species Figure Holotype. Fingers one-half webbed. their distance from end of snout about one-third that from eye. opposite side from "La Macarena. Snout moderate. tongue three-fourths as wide as mouth opening.12. Cabrera. — USNM 20.12.5. palpebral membrane not reticulate. Rio Payamino. Where the cotypes are is pale to medium brown. Paitanga. however. Hyla cabrerai. collected along the Catio Guacaya. 1. — CJG 2317. Eye moderate.277 (cotype) MCZ Maranon.. 80. Cabrera in collected along the Cano Tui between Mitu and the Raudal de Yurupari.184-9 Meta: Upper Rio (cotypes).12. Vaupes. it is plain to see pale yellow. from Amapa Territory. CNHM 54765.215-7 (coPastaza River between Canelos and 19695-6. about three-fourths the diameter of eye. Colombia. the CNHM specimen cotypes look as though they didn't have quite enough. A moderate-sized Hyla with a patagium. Amazonas. Description oj holotype. oblong choanae. the latter apparently was preserved in too strong formalin. BM 80. cabrerai is most closely allied to H. and the fingers about one-half webbed. separated from each other by an interval equal to about two-thirds their distance from eye. Paratypes.238 types). throat and chest heavily sprinkled with small but distinct dark brown spots. Vomerine teeth in two moderate. considerably projecting. Tympanum very distinct.5. a tributary of the lower Rio Apaporis. ECUADOR: Alto Napo. nostrils more lateral than superior.000 m. and where the cotypes are brown the Chicago Natural History Museum specimen is nearly black.5. interorbital distance slightly greater than width of upper eyelid.FROGS OF COLOMBIA —COCHRAN AND GOIN 215 that the same basic pattern obtains. a distinct pattern on the back. 80. not particularly prominent. buckleyi. from which it differs in having chest and throat spotted and a greater amount of webbing between the fingers. Plate 29a-c 152759. and WCAB 13284. broadly ovate. its diameter nearly equal to its distance from nostril. May 1952. . (two cotypes).12. fourth finger consider- — — . while the fully. Canelos. truncate in profile. separated from eye by a distance nearly equal to onehalf its own diameter. slightly arched series." USNM 152199.5. Canthus rostralis well defined loreal region concave and oblique. H. WCAB 294. BM 80. Specimens Examined COLOMBIA Giiejar. its posterior border but slightly free and unnotched.

See also Plate 29 a-c. Body somewhat elongate.third the tympanic area. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. . traces of a Figure 20. a row of conical. when hind leg is adpressed. USNM 152759 (same size. no projecting rudiment of a pollex.S. in postaxillary region a little narrower than greatest width of head. which are X 1. soft. with wart-like tubercles arising from various areas in which the glandules of the skin appear particularly concentrated. on upper eyelids and occiput. holotype. profile Hyla cabrerai. when hind legs are bent at right angles to body.3). a distinct narrow glandular ridge encircling upper part of tympanum. ulnar ridge present but somewhat indistinct. knee reaches nearly to axilla.— 216 U. reaching to middle of disk of third which covers about one-half the tympanic area. heel reaches tip of snout. heels overlap greatly. that of belly and lower surface of thigh uniformly and very finely granular. the web on fourth toe reaching base of the disk. these tubercles particularly conspicuous on snout. and around anus. A small patagium extends from the back of upper arm to side of body. when limbs are laid along the side. NATIONAL MUSEUM BULLETIN 2 88 ably longer than second. third and fifth toes subequal. a small dermal appendage on heel. Skin of upper parts glandular. skin of throat and chest nearly smooth. wartlike structures along the tarsal ridge. except head and mouthparts. disk of fourth toe covering about one. Toes webbed. including loreal and canthal regions.

posteriorly diverging lines in the suprascapular region. — Dimensions. posterior projecting portion that passes back to the occipital region. narrowly separated crossbands on each thigh. a line along the supratympanic ridges. each chin. discrete —The three specimens at hand (holotype and two paraand pattern. Color in alcohol. the other two species have the webbing reduced closely knit group.5 mm. hand. 19. his botanical and of of his zoological collections.2 mm. and a transverse glandular ridge above the vent. a somewhat enlarged blotch on each sacral protuberance that extends to the groin on each side. tibia. together with buckleyi and pearsoni certainly make a We have examined the type series of buckleyi in detail and have seen (but have not examined in detail) the type of pearsoni. a pair of straight. between the fingers. 38. a single broad crossband on each upper arm. types) are rather uniform in both structure insignificant... shank. and breast with numerous about the diameter of a pin head. have long hind legs. his native country by both 8. while it is heavily spotted in cabrerari and is reticulated in pearsoni. In addition to H.. R. throat. buckleyi has the belly dusky.. H. cabrerai seems to have the heaviest hands with the most webbing between the fingers.1 mm. a triangular dermal appendage on the heel. frog. while in pearsoni the extended heel reaches only to the eye. femur.FROGS OF COLOMBIA — COCHRAN AND GOLN 217 skinfold across chest. The specimen from Amapa. and foot. three broad. The species that are known from life have pink or or red webs between the fingers and toes. Isadore Cabrera. no vocal sac. This species is dedicated to our friend Sr. a blotch on each can thai region. albo- . H. Both buckleyi and cabrerai. a narrow interocular bar with a median.. 31. and hand. 52. This species. has a little less webbing between the fingers than the other two but the difference is so slight that it seems Remarks. with the extended heel reaching to the tip of the snout. heel-to-toe. head width.2 mm. E. distinct.. 27. roof of skull not exostosed. spots. collector who was trained in the field by Prof. ALBOMARCINATA group Moderate-sized. The —A rather brilliantly marked black and yellow general ground color seems to be a pale yellow with a black pattern as follows: a few tiny blotches along margin of upper lip. adult female.8 mm. 19.7 mm. Skin of head not coossified with skull. pale tree frogs (pale to white in preservative) with patagia and well-developed ulnar and tarsal ridges. Schultes Harvard and who has contributed appreciably to the knowledge of the type. Brazil. Head and body. 17. forearm. head length.2 mm.

164. 98. 97. p. 1843. 1930.— Giinther. p. 1871. 1-6. 555. 140.— Peters.— F. p. 273. 1930. p. pi. 1931. p. fig. a2 . pp.— Cope. vomerine teeth in two : moderate-sized. p. vomerine teeth in two H. 533. — Phyllobius albomarginatus. p. — 1860. 4. p. 1867. Ecga [Ega]. 284. 35. 38. p.— Beebe. its posterior border fused and somewhat truncate . p. 2 (separate). 132. Lutz. 57. 33. CJG 2354. p. 1919. 1860. Colombia. life). 1927. 13. 43. from which it strongly developed vomerine tympanum and more teeth. 555. p. p. albpunctulata small transverse series Tympanum about one-half the diameter of eye. pi. Hyla massarti De Witte. p. 162.— Cunningham. p.S. 1925. from Zetaquira. 143. 1882a.. a small. p. 1935. — — — 1824. tongue three-fifths as wide as mouth opening. 1926. p. 771. 1872a. 1897. 101. H. 56. 1930. 1907. albopunctulata. p. Hyla cinerascens Spix. 124 Nieden. The two species known from Colombia may be separated by the following key: a1 .— Reinhardt. Lutz. 13. 174. 1860. 1900. 1858. fig. 1830. 1838. 1961. p. 511. Wagler. 1923. 1825.— Wandolleck. p. p. p. — Description. Phyllobius exanthematicus Fitzinger. somewhat arched . 1926. Botanical Gardens. 1917. 33.— Boulenger. p. 31. p. 1824. p. Tschudi. Tympanum about one-fourth the diameter of eye.— Werner. 1841. 207. 218 U. p. 1830. pi.— A. 468. Fitzinger. Hyla infulala Wied. fig. p.— Baumann. albomarginata series Hyla albomarginata Spix Plate 29f-h 1824. 1928. albopunctulata of Colombia. 1955. pp. 413. p. near Miraflores. 30. 1912. 1886. lying close together just slightly behind the posterior borders of the moderatesized. 225 (type locality. H. Bahia). 595.— Burmeister. 298. rounded choanae.— 72. pp. on the Solimoes.— Crawford. 1882a. Rio de Janeiro) —A Hyla with ulnar and pure white in feet in The tarsal ridges. 671 (type locality not given). p. NATIONAL MUSEUM BULLETIN 2 8S marginata and H. 201. p. 30. p. Hyla albomarginata Spix. 1946. p.— Fouquette. city of Diagnosis. 4 (type locality. 2 Cochran. fig. — Fitzinger. 356. 8. p.— B. 1 (type locality. figs. triangular and practically no dorsal pattern (nearly preservative but with bright pink webbing on hands and dermal appendage on heel. p. rujitela of Panama also is a member of this group. Midler. 186 Steindachner.— Mertens. p.— Myers. 1950.— Parker. 1931.— Ahl. 8. and Lutken 1862. 260. Hypsiboas albomarginatus. 1901. broadly rounded. 412. 1856. p. Boyaca.— Dumeril and Bibron. 220. 1949. p. albomerginata apart tarsal ridges from all Colombian differs in its larger frogs except H. p. Vomerine teeth in two somewhat arched series. nearly pure white dorsum in combination with ulnar and and dermal heel appendages set H. 74.— Miranda-Ribeiro. p.

palpebral membrane not reticulate.FROGS OF COLOMBIA — COCHRAN AND GOIN 219 but unnotched. the by a distance nearly equal to its own diameter. adult female. when limbs are laid along the side. 13. Eye not large.5 mm. fourth finger longer than second. femur. A small patagium extends from the back of the upper arm to the side of the body. inconspicuous.. 168). when hind leg is adpressed. Dimensions. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. not projecting. separated from eye slightly sloping in profile. the upper lip flaring out slightly below it. knee and elbow overlap slightly. no apparent traces of a skinfold across chest.. in postaxillary region a little narrower than greatest width of head. Although H. a very narrow black line margins the upper lip back to a level between the naris and eye on each side. a small but distinct triangular dermal appendage on heel. Skin of upper parts smooth. which is about equal the distance between nostrils. no projecting rudiment of a pollex.0 mm. Remarks. interorbital distance half again the width of upper eyelid. Canthus rostralis slightly defined. Toes nearly completely webbed.6 mm. heels overlap greatly. 21. Snout moderate. Body rather elongate. Fingers webbed at base.. but prominent. and dermal appendages on heels are milky white above and black below. rounded when viewed from above. that of belly and lower surface of thigh uniformly and very finely granular. a distinct ulnar ridge. hand. the web on fourth toe reaching the middle of the penultimate phalanx.4 mm... a distinct tarsal ridge. nostrils more lateral than superior and more anterior than lateral. Skin of head not co-ossified with skull. when hind legs are bent at right angles to body. head width. tiny black spots on dorsum and top of head. the ulnar ridges are milky white with interrupted black lines. skin of throat and chest smooth. heel reaches almost to nostril. Color in alcohol. A transverse dermal fold above vent. just reaching to disk of third which about covers the tympanic area. 1955 p. 12. tarsal ridges. Tympanum distinct but not very prominent. each naris margined in black. no vocal sac. heel-to-toe. its diameter equal to its distance from nostril. Head and body. 40. disk of fourth toe covering about two-thirds the tympanic area. albomarginata has been previously recorded from Colombia (Cochran. tibia. separated from each other by an interval equal to about their distance from eye. head length. 28. 13. upper jaw hardly extending beyond lower.2 mm. loreal region somewhat concave and oblique. the transverse fold above the vent. 19. third and fifth toes subequal. 600) pointed out that the specimens on which the Colombian records are — — — .. roof of skull not exostosed. Practically white with a pattern of black pigment as follows: scattered. their distance from end of snout about onehalf that from eye.5 mm.7 mm. Fouquette (1961 p. about three-fifths the diameter of eye.

it tympanum and 1908. and albomarginata a representative east of the Andes. tongue two-thirds as wide as mouth opening. by having ulnar and respectively. 385 (type locality. 21. MZUM 104146 (3). 271. MZUM 104143 (10). near Miraflores.2. its posterior border fused and very shallowly notched. Hyla palmeri Boulenger. Jimenez). Ecuador. There seems little doubt that albopunctulata is a representative of this group west of the Andes. 1964a. from which smaller vomerine teeth. p.32] (a cotype of palmeri). the upper jaw extending slightly beyond lower. The specimen described above differs from the examples from eastern Brazil in the amount of webbing on the hands and amount of dorsal pigmentation. 264. F. Barro Branco. wider than long. separated from each other by an interval about equal to their distance from eye. loreal region slightly concave and somewhat oblique. southwestern Colombia.— Gorham.— BM Hyla albomarginata. CJG 235. and it may be that when adequate material becomes available it will prove to be distinct from albomarginata. 515 (type locality. Can thus rostralis slightly defined. p. truncate in profile. albomarginata of eastern South America. rufitela a representative in Panama.13. NATIONAL MUSEUM BULLETIN 288 based were small. Valle. 303. Vomerine teeth in two small. p. D. Colombia. Hyla albopunctulata Boulenger Plate 29d. Specimens Examined COLOMBIA Boyaca: Zetaquira. p. Perhaps differs it is by its most closely allied to smaller Description. ruiitela of Panama or H. snout moderate.220 U. from Jimenez. 1963. as defined by him.68 [1947. 104145. Diagnosis.5.— Gorham. Re-examination of the specimens mentioned above convinces us that they should not be identified as albomarginata and that the only unquestioned Colombian record is the specimen described above from Boyaca. Hyla albopunctulata Boulenger. poorly preserved specimens that are not conspecific with either H. rounded choanae.e 1882. somewhat rounded when viewed from above.— Rivero. 1963.— Nieden. nostril more lateral than superior. Sarayacu). 1923. BRAZIL: (2). 1923. tarsal folds along the verse vomerine series lying slightly behind the small rounded choanae. broadly cordiform. transverse series lying close together and just behind a line connecting posterior margins of the small.— Nieden.. 1908. 21. p. the upper lip . 104144 Rio de Janeiro. p. slightly projecting. 1882a.4 Rio de Janeiro. terized —A somewhat granular Hyla characarms and feet a wavy glandular fold over the vent. p. and a short transmoderate-sized.S. their distance from end of snout about one-half that from eye.29.

1 mm. disk of fourth toe covering the tympanic area. that of belly and lower thigh uniformly and finely granular.12. Body not elongate.230) and another one (80. when hind legs are bent at right angles to body. which makes contact with that on leg when limbs are folded. is well-marked with scattered. its diameter than its distance from nostril. a quite distinct but rather narrow glandular ridge encircling upper part of tympanum.12. the cotype of albo- (BM 80.. In all the speci- punctulata from Sarayacu 159) . the web on fourth toe reaching base of the penultimate phalanx.5. no projecting rudiment of a pollex.. knee and elbow touch. traces of a skinfold across the chest. 29.. tibia.5 mm. Toes about three-fourths webbed.. which is relatively wide.5 mm.29. anterior to groin. fourth finger but slightly longer than second.FROGS OF COLOMBIA —COCHRAN AND GOIN 221 it. foot. Eye moderate. hand. just reaching to disk of third which is half again as broad as the tympanic area. head length. Although the large cotype of palmeri is firm and exceedingly well preserved while the cotypes of albopunctulata are quite soft Color in alcohol. heels overlap slightly. 20. — Dimensions. when hind leg is adpressed. 14 mm. Fingers one-third webbed. head width.7 mm. interorbital distance half again as wide as upper eyelid. brownish. not prominent. 13. Head and body. when limbs are laid along the side.13.. metatarsal tubercles not distinct. roof of skull not exostosed. a well-developed fold from the elbow along the ulnar side of the arm to the wrist. and about equivalent to distance between nostrils. 13. tarsal fold somewhat whitish in color. clear white spots about the size of a pencil dot. including that of the limbs. — A faded yellow all over except for scattered. — and flabby. Remarks. 41. no apparent vocal sac. pepper-like markings on middle of back. a very distinct tarsal fold continuing to the heel where it terminates in a small but distinct triangular appendage. Skin of head not co-ossified with skull. Tympanum distinct. heel reaches between eye and nostril. 21.. small. about one-fourth the diameter of eye. third and fifth toes subequal.5. Skin of upper parts very finely granular.32-3]). for in one of the cotypes of albopunctulata the entire dorsal surface. for the it is name is evident that they all are the same species.from "Ecuador" lack these white punctulations. a similar fold above vent. no inguinal gland. femur.68-9 [1947. A few white spots can be seen on two of the other cotypes.2. but the cotypes of palmeri (BM 1908. skin of throat and chest finely granular. The reason evident. separated from eye by a dis- flaring somewhat below slightly greater tance nearly equal to its own diameter. A patagium extends from the back of the upper arm to the side of the body. in postaxillary region about equal to greatest width of head.5 mm. and on each side. is a little fold which seems to be a structural feature.5.8 mm.

105. 207. Key a . 1933. 1830. 1843. p. Hyla punctata. Wagler. spots or flecks on the dorsum. H.— Miranda-Ribeiro. p.— B. 80. 207.13. p. 201. 13. p. p.29. p. 80. p. 403. 48.— Daudin. p. p. p.— Baumann. p. 1829.5. 1802. 1952. 101.— Gravenhorst.—Gorham. 662. 1963. 357. 1 b2 a2 Colombian Species in the punctata Group Red 1 ft to . 1856. bl . Calamita punctata Schneider.. p. 201. Valle: BM ECUADOR: . punctata only about three-fifths the diameter of eye Dorsum without distinct red markings. and without ulnar or tarsal ridges. B. 23. tympanum H.5. Hyla granosa gracilis Melin.S. Hyla variolosa Spix. 208. Gunther. 1937a. p. without spines on the pollex.— Goin. Magdalena. p. p.40— Parker..5. — — — 1824. 1958. 1802. — — Wagler. 1896. tympanum about H.— Peracca. p. p. 41. p. Vomerine teeth in two somewhat arched series Hyla punctata (Schneider) Plate 24d-f 170 (type locality. 1871. p. Colombia. rubeola two-thirds the diameter of eye Dorsal spots discrete rounded spots rather than minute flecks. p. 1843. 201. p. 1912. p. 1964a.— Boettger. p. 30. 1919. p. 303. — p.— Cope.— Peters. 56. Hypsiboas punctatus— Fitzinger. BM 9. 1889. 1926. p. p.— Nieden. 101.— Boulenger. p. 1951a. b2 . 301. what little brown pigment is present tends to be concentrated anteriorly on the back and on top of the head. p. 81. 1923. p. . Goin and Layne. Specimens Examined COLOMBIA BM 1908. . Dorsal spots minute flecks rather than rounded spots.12. 1892. 1872.2. p. 1858. 1872a. 126. 1874b. 88. 37 (type locality. 90. Lutz. 163.12. 174. 30.— Berg. 1951a. 554. — Lutz. Fundaci6n). 247. 1922. PUNCTATA group Moderately small green tree frogs with small vomerine tooth patches. Crawford and Jones. NATIONAL MUSEUM BULLETIN 2 88 mens. 1841. p. pp. Jimenez. 552. 1935. Scinax variolosa. 1871a. p. 1904. Auletris variolosa. p. 1933. 307. 34. 21. p.— Rivero. but four now known from Colombia may be distinguished with the aid of the key. "America"). Hyla granosa. 120.230 (cotype of albopunctulata) no particular Sarayacu. p. 1958. 771.159-62 (cotypes of albopunctulata) locality. It is not now known just how many South American species are in this group. Santa Marta Mts. 222 U. 1882a. 40.— Beebe. p. 1880. 121. 1867. p. Lutz. p. 1941.— A. 1830.32-3] (cotypes of palmeri). 303. 4. 1830. 1927.— . 1799. reduced webbing between the fingers. 1898a. Hylella pearsei Ruthven. 1873a. p. 1803. rounded patches Hyla sp. p. . Dumeril and Bibron. woods along Amazon River). rhodoporus Vomerine teeth in two small.— Liu.68-9 [1947. 1933. 1830. 37.Burmeister. 57 (type locality. p.

from Leticia. parts smooth. part of No when hind leg is ad- are laid along the side. Toes slightly less than onehalf webbed. interorbital distance. loreal region a little concave and somewhat oblique.A small "green" frog with greatly reduced webbing between the fingers and with small patches of vomerine teeth. fourth than second. indistinct. rounded choanae. low. Eye not large. Colombia. upper eyelid. rounded outer metatarsal tubercle. pressed. and distance its skin. Amazonas. legs are bent at right patagium. nostrils more lateral than superior. heel reaches tip of snout. somewhat projecting. rounded patches. when limbs when hind angles to body. Description. the web on fourth toe reaching to middle of antepenultimate phalanx on the lateral side and to its base on the medial side. Vomerine teeth in two short.FROGS OF COLOMBIA — COCHRAN AND GOIN 223 — Diagnosis. Canthus rostralis fairly well defined. broadly cordiform. in postaxillary region a little narrower than greatest width of head. tongue two-thirds as wide as mouth opening. the upper jaw extending but little beyond lower. lying close together between the moderate-sized. their distance from end of snout about one-third that from eye. no projecting rudiment of a pollex. heels do not quite meet. a distinct oval inner and a smaller. Fingers with but a trace of web at base. — black choroid coat standing out sharply against the pale yellow from nostril. U-shaped when viewed from above. knee and elbow slightly overlap. disk of fourth toe covering about onethird the tympanic area. its diameter equal to its distance between nostrils about equal. palpebral membrane not reticulate. Snout moderate. just reaching to disk of third finger considerably longer which covers about two-thirds the tympanic area. Tympanum the diameter of eye. separated from each other by an interval equal to about two-thirds their distance from eye. Hyla punctata may be distinguished from rubeola by having rounded red spots (rather than tiny flecks of red) on the back and by its more sturdy build. its posterior border slightly free and shallowly notched. vocal sac not . skin of throat and chest very finely granular. but prominent. no dermal appendage on heel. truncate in profile.. traces of a skinfold across chest. by a about three-fifths distance equal to one- diameter. Skin of upper a rather narrow glandular ridge encircling upper tympanum. separated from eye half its own distinct. a low ulnar ridge. UF 8518-1. that of belly and lower surface of thigh uniformly and not so finely granular. third and fifth toes subequal. It may be distinguished from rhodoporus by having the tympanum equal to about three-fifths (rather than two-thirds to three-fourths) the diameter of eye and by the presence of rounded red spots on the dorsum. Body not elongate. adult male. the upper lip flaring out moderately below it. no tarsal ridge.

224 NATIONAL MUSEUM BULLETIN 288 U. punctata. 12. MZUM 54639 (type of Hylella pearsei). As Dr. 20. —A yellowish ground color with rather distinct The dorsal white spots. Head and body. Some specimens have no apparent spots on the dorsum.5 mm. and the series from Trinidad are no more than one would expect between two series collected and preserved by two different collectors at two different times. Some individuals at night are nearly a uniform rusty red dorsally. Variation.. where there seem to be a number of distinct but closely related forms and an even greater number of available names. granosa 10987-8. .9 mm. apparent. 35. 467 (type TRINIDAD: UF NM 3647.. at daylight. posterior to — — Specimens Examined COLOMBIA Amazonas: Magdalena: Leticia. about one scattered white spots dorsally. This difference surely is due in part to the time the various specimens have been preserved. Upper Rio Vaupds.9 mm. the belly being more brownish yellow than the rest. Skin of head not co-ossified with skull. hand. and. 1951a). Lutz points out (B. the species of allied "green Hylas" are a "difficult group. they are conspecific. if kept alive. will have faded to greenish brown dorsally with distinct yellowish spots.7 mm. 7300 (8). 41-7893 ex. mm. 12. others are quite distinctly spotted.. head length. Fundaci6n.. Nevada de Santa Marta. gracilis). The differences between the series from Leticia in Amazonas. The series of fresh specimens at hand from Leticia were compared with a cotype of H. H. 10. Colombia. head width. as far as we could determine from the preserved specimens. are scattered over the top of head and dorsum.S. 8519-20. granosa gracilis Melin.9 Color in alcohol. The ventral surfaces are yellow. A very narrow. Remarks. punctata. and the same individuals. 18.5 mm. indistinct white stripe originates on each side just tympanum and terminates just anterior to groin. — Dimensions. heel-to-toe. The main difference discernible in any series of specimens of punctata that we have seen is in the degree of spotting on the back. Lutz. roof of skull not exostosed. but it may be due in part also to the hour of day when the specimens were preserved. USNM 151944-5. 10986 (2)." and as yet no one has had the opportunity to study all of the group's various forms in life. . millimeter in diameter. We also examined the type of Hylella pearsei Ruthven (MZUM 54639). Meta: BRAZIL: Near Villavicencio. seems to be rather uniform over a rather extensive range. 7727 (2). and it appears to be an immature specimen of H. Sierra UF 8517. 25. of H. as now understood. tibia. This is particularly true in the northern Amazon basin. femur.. 8518 (6).5 mm.

disk of fourth toe covering about one-sixth the tympanic area. and a more slender build. no projecting rudiment of a pollex. rhodoporus it differs in its more slender build and the presence of dorsolateral is allied to punctata and rhodoporus. Colombia. loreal region somewhat concave and oblique. Fingers with but a trace of web at base. from above. when limbs are laid along the side. about three-fifths the diameter of eye. Medem. greatly reduced webbing on the ringers. the tympanum separated from the eye by a distance equal to only one fourth its own diameter. third and fifth toes subequal. January 4. the web on fourth toe reaching base of the antepenultimate phalanx. Vomerine teeth in two series. It is characterized red stripes and the tiny fleck-like red specks on the back. From H. tongue one- mouth its posterior border U-shaped when viewed all free. when hind legs are bent at right angles to body. narrow. no dermal appendage on heel.FROGS OF COLOMBIA —COCHRAN AND GOIN 225 Hyla rubeola. which is slightly greater than distance between nostrils. the upper jaw extending but slightly half as wide as opening. truncate and not at beyond lower. interorbital distance than width of upper eyelid. knee and elbow considerably overlap. Tympanum very distinct. just reaching to disk of third which covers about one-fourth the tympanic area. no ulnar ridge. — Description of holotype. Canthus rostralis strongly defined. not particularly prominent. Toes less than one-half webbed. Skin of upper parts smooth. truncate in profile. and (iD life) bright red. Body elongate. heel reaches slightly beyond tip of snout. Meta. dorsolateral stripes and scattered pepper-like spots of bright red on the back. a rather narrow glandular ridge passing nostril. nostrils nearly lateral. the upper lip flaring out below it. fourth finger considerably longer than second. heels fail to meet. their distance from end of snout about one-fourth that from eye. palpebral slightly greater . a distinct oval inner and an equally distinct rounded outer metatarsal tubercle. new species Plate 30a-c — USNM 152751. —This new Holotype. It may be differentiated from punctata by having the red dorsal markings as minute red flecks rather than as distinct rounded spots. 1959. Eye moderate. no tarsal ridge. broadly triangular. collected on Serrania de La Macarena. considerably projecting. oblong choanae. lying close together between the moderate-sized. Snout moderate. by F. by having a small tympanum set close to the eye. separated from each other by an interval equal to about one-half their distance from eye. separated from eye by a distance nearly equal to one-fourth its own diameter. No patagium. a member the "green" Hyla group. in postaxillary region distinctly narrower than greatest width of head. its diameter equal to its distance from membrane not reticulate. Diagnosis. of species. when hind leg is adpressed.

head length. A distinct but quite of small dorsally pattern originates on each side above tympanum and stripe red narrow — passes posteriorly to region of the sacral hump where it fades out. Hyla rhodoporus Giinther Plate 30d-f 1869. hand. Remarks. its posterior —CNHM to . This frog seems to be another of the species endemic of — Serrania de La Macarena. p. by having the tympanum equal about three-fourths (rather than one-half to three-fifths) the diameter of the eye.. Skin of head not co-ossified with skull.8 mm. Description. greatly reduced webbing. although careful inspection will demonstrate its definitive characters of a small tympanum and the We did not see the specimen alive but while still fresh it was spectacularly colored with its brilliant red dorsolateral stripes and with the dorsal surfaces flecked with bright red as though someone had sprinkled it with cayenne pepper. lying close together between the moderate-sized. 307. two on each forearm. with small patches of vomerine teeth. — Dimensions. three on right thigh. Meta. 1923. 662. 1952. 33 mm. Head and body.. NATIONAL MUSEUM BULLETIN 288 above upper part of tympanum. locality. tongue one-half as wide as mouth opening. H. no vocal sac. 1872. and sixteen scattered over the back.— Lutz and Kloss.— Nieden. no traces of a skinfold across chest. 357. five on left shank. p. 313. head width..— Peters.4 mm. 9. —A small "green" Hyla with greatly reduced webbing between the fingers. roof of skull not exostosed. 81786. femur. 17.6 mm. Upper Amazons). p. skin of throat and chest smooth. p..S. p. Except for these red markings. heel-to-toe. somewhat rounded patches. Lutz.226 U. two on left thigh. pale yellow. p. broader than long.— B.. and shanks are scattered tiny but distinct cardinal red spots. 17. After several years in preservative the type has faded to a pallid yellow and might be taken at a glance for some of the other "green" Hyla from northern South America. tibia.— Boulenger. Hyla rhodoporus Giinther.9 mm. thighs. rhodoporus may be distinguished from rubeola by its sturdier build and the absence of dorsolateral stripes. 1871.. that of belly and lower surface of thigh uniformly and very finely granular.4 mm. 1882a. four on right shank.7 mm. 10. 21. are two such spots the entire specimen is clear. Villavicencio. 1951a. 10. On top of head and back and on the dorsal surfaces of forearms. subadult female. There on top of head. 403. It differs from punctata Diagnosis. Vomerine teeth iu two short. red spots and stripes. Colombia. 653. rather oblong choanae. 480 (type p. Clear pale yellow above and below with a distinct Color in alcohol.

p. Skin of head not co-ossified with skull.third that from eye..FROGS OF COLOMBIA — COCHRAN AND GOIN 227 border fused and unnotched. Lutz (1949. Can thus rostralis rather well defined. disk of fourth toe covering about one-half the tympanic area. head width.. No patagium. a narrow glandular ridge passing above tympanum. fourth finger but little longer than second.. rhodoporus was considered a synonym of II. 10 mm. rhodoporus . width of upper eyelid. H. 13. just a hint of a skinfold across chest. and distance between nostrils about equal. the upper jaw extending but little beyond lower. nostrils more lateral than superior. — Remarks. somewhat transluscent upper Color in alcohol. Toes less than one-half webbed. When hind legs are bent at right angles to body. heels just snugly touch. interorbital distance. about four-fifths the diameter of eye. somewhat projecting. 13 mm. palpebral membrane not reticulate. head length. tibia. not quite reaching to disk of third which covers about two-thirds of tympanic area. no dermal appendage on heel. Geographically. heel-to-toe. in postaxillary region a little narrower than greatest width of head. U-shaped when viewed from above. 313) pointed out in her synopsis of the "green" Hyla that it probably was a valid species. femur. —This specimen is a uniform yellowish tan above and below without vestige of a pattern. punctata until B. knee and elbow considerably overlap. no ulnar ridge. hand. 18. — Dimensions. 18. separated from each other by an interval equal to about twothirds their distance from eye. Body not elongate. Snout not large. Eye not large but prominent. the upper lip flaring out appreciably below it. Tympanum very distinct.. a distinct oval inner and a smaller rounded outer metatarsal tubercle. that of belly and lower surface of thigh more coarsely granular. adult male. 26. Skin of upper parts very finely glandular. no projecting rudiment of a pollex.3 mm.. the web on fourth toe reaching base of the penultimate phalanx. when limbs are laid along the side. its diameter equal to its distance from nostril.5 mm. The dark choroid coat of the eye shows through the yellowish. skin of throat and chest finely granular. loreal region slightly concave and rather oblique.3 mm..8 mm. to — The only variations discernible in the series at hand be results of preservation and the ages of the various Variation. no tarsal ridge. 37. seem specimens. Head and body. vocal sac subgular. Fingers webbed at base. roof of skull not exostosed. when hind leg is adpressed. their distance from end of snout about one. heel reaches tip of snout.8 mm. separated from eye by a distance equal to onefourth its own diameter. rather truncate in profile. eyelids. web of third finger reaching base of antepenultimate phalanx. third and fifth toes subequal.

of the 2 88 Andes. GRANOSA group Medium-sized green Hyla in which the male bears a protruding spine on the pollex. 117511) from Rio Guaimaral. furthermore. The it little more dried-out than the described specimen. while the tympanum of punctata is only about one-half the diameter of the eye). 10. Specimens Examined COLOMBIA Caqueta: Florencia. its tympanum is relatively larger than that of albomarginata as redefined by Fouquette (1961. able.2. with the vomerine teeth in rather heavy. Hyla A single specimen (USNM we prefer to leave without a seem to align it sp. 595) it also differs from that species in lacking a super-anal white line. the species rhodoporus is seen to differ in having a broader head and more rounded snout. 66. USNM 152069-71. in having a larger tympanum (two-thirds to three-fourths the diameter of the eye. traces of light dots are still discernible on the back of the type seems to be a for its toes Very faint type. USNM 152112-4.23. and a white ulnar ridge and white line on the tarsus.5] (type of rhodoporus). Its reduced webbing and build with the "green" Hyla group. and in this respect it is less similar to albomarginata than it is to the other species mentioned above. Putamayo: Puerto Asfs. 60444. 152230-46. and the webbing between them are a little more shrunken. it has greatly reduced webbing on the fingers.9. On the other hand.S. When compared directly with punctata. Since the specimen is faded and has been preserved for quite some time. When the type the above-described specimen was directly compared with was found to differ only in being a little more robust. and in being less conspicuously spotted. arched . and the fact that its vomerine tooth rows are somewhat arched would seem to indicate that it might be allied to that species. name.4 [1947. white bars on the heels. and rhodoporus. CNHM 81785-6. Atlantico. 81783. Meta: Villavicencio. rubeola. while punctata possibility exists that rhodoporus may- represent a race of punctata. p. we prefer to hold specific identification in abeyance until fresh material from the north coast of Colombia becomes availferent from the three species of that group presently recognized Colombia punctata. NATIONAL MUSEUM BULLETIN seems to be a species of the eastern slopes is The a species of the lowlands. but it is certainly dif- from seems more like albomarginata.26. In general it . USNM "UPPER AMAZONS": BM 152280-1.— 228 U.

July J.— FROGS OF COLOMBIA The series. A H. broadly cordi- Figure 21. while the lack of heel appendages distinguishes it from H. rubracyla H. Paratype — CNHM 81922. lying very close together between the rather large. without dermal appendages on the heels. Dorsolateral stripes present b2 No . . new species Figure 21 Holotype. . three species —COCHRAN AND 229 GOEST now known from Colombia may be differenti- ated by the following key: a No 1 a2 . 12. guibei dorsolateral stripes triangular dermal appendage on the heels Hyla rubracyla. arched \ series. Choc6. granosa H. Colombia. Diagnosis. — / Description of holotype. The presence of dorsolateral stripes serves to differentiate this new species from H. — USNM 157820. Valle. 1961. Colombia. by Fred Medem and Coleman Goin. near Cdrdoba. 337-262—70 Hyla rubracyla. bl . dermal appendage on heel. collected along the Rio Calima. —A medium-sized green Hyla without a transverse white ridge above the vent. guibei. granosa. collected at Aqua Clara. and with conspicuous dark canthal and dorsolateral stripes. holotype. Vomerine teeth in two heavy. tongue one-half as wide as mouth opening. oblong choanae. 16 USNM 157820 (same size).

roof of when hind No skull not exostosed.5 Color in alcohol. nostrils lateral. A conspicuous. Canthus rostralis strongly defined. very dark reddish brown bar between the nostrils which continues back along the canthus rostralis and along the outer margin of upper eyelid. considerably projecting. 37. and a much smaller dark spot on dorsal surface of each thigh just above knee and one across dorsal surface of forearm. 19. — Ground color white. rounded when viewed from above.. truncate in pro- file. its diameter equal to its distance from nostril. Head and body. when limbs are laid along the side. its NATIONAL MUSEUM BULLETIN 288 posterior border very slightly free and shallowly notched.. the upper jaw extending a little beyond lower. about two-thirds the diameter of eye. a low ulnar ridge. no transverse glandular white ridge above vent. the web on fourth toe reaching base of the penultimate phalanx. Toes slightly more than one-half webbed. no traces of a skinfold across chest. patagium. reaching to disk of third which covers about two-thirds the tympanic area. which is slightly greater than distance between nostrils. Eye not particularly large but rather prominent. Skin of head not co-ossified with skull. no dermal appendage on heel. hand. 28.7 mm. Skin of upper parts very smooth. that of belly and lower surface of thigh uniformly and a little more coarsely granular. palpebral membrane not reticulate. mm. Body not elongate. when hind leg is adpressed. form.230 U. separated from each other by an interval equal to about two-thirds their distance from eye. femur. Tympanum very distinct. interorbital distance slightly greater than width of upper eyelid. 50. Elsewhere the dorsal surfaces are peppered with little rounded specks. tibia. separated from by a distance equal to one-third of its own diameter.S. head width. these being thickest on top of head and dorsum. 18.. loreal region concave and oblique. Interrupted. — Dimensions. heels overlap greatly. fourth finger longer than second by the diameter of eye a disk. a distinct oval inner and a smaller indistinct rounded outer metatarsal tubercle. knee and elbow overlap considerably.. a spiny projecting pollex. Each shank marked above with three distinct narrow. in postaxillary region a little narrower than greatest width of head. third and fifth toes subequal. Adult or subadult male.4 mm. skin of throat and chest very finely granular. disk of fourth toe covering about one-half the tympanic area. heel reaches beyond tip of snout. less . a rather narrow glandular ridge passing above tympanum. legs are bent at right angles to body.1 mm. an interocular dark spot. less-conspicuous dorsolateral stripes of the same color. 28. dark spots. no tarsal ridge. the upper lip flaring out below it.. heel-to-toe. their distance from end of snout about onethird that from eye.7 mm.2 mm..8 mm. Snout moderate. Fingers webbed at base. 15. no vocal sac apparent. head length.

the type was conspicuously life. p.— Nieden. 220. The name refers to the bright red markings on the face. Except for having the pepper-like markings tinier and more sparsely distributed and lacking the little dark marks on each thigh just above the knee. Snout large.— Beebe. and without dorsolateral dark stripes. Caqueta. 291. the next morning. 301.— Gorham. here restricted. The absence of heel appendages differentiates H. 1919. 99. sitting Hyla granosa Boulenger Plate 31a-c 1S82. does not penetrate the skin as does that of the male. 33. slightly rounded in profile. orange-tinted yellow. 1897. without a glandular ridge above the vent. and the absence of dorsolateral stripes distinguishes it from H. 1882a. Demerar a Falls). and on top of the two outer fingers. 358 (type locality. Vomerine teeth in two / ~\ -shaped series. 1963.— Werner. broadly U-shaped when viewed from above.FROGS OF COLOMBIA — COCHRAN AND GOIN 231 and along sides below dorsolateral dark stripes.— Noble. p. its posterior border fused and shallowly notched. rubracyla. Description. on outer side of hand. 208. her markings are identical with those of the type. p. Colombia. p. British Guiana. tongue two-thirds as wide as mouth opening. nostrils more lateral than superior. the upper jaw extending but little beyond lower. In daylight.— Lutz. — Remarks. somewhat elongate choanae. from Florencia. guibei. The voice was not heard. 1931. 1961. in in the type.— Crawford. The type was collected at night. with the pattern in dark red standing out prominently. while palpable. 1923. —A pale frog without dermal heel appendages. p. 30. along outer margin of foot. along the top of the shank. Hyla granosa Boulenger. 22. USNM 152756. p.4 millimeters head-body length. 1951^. p. broadly cordiform. and the canthal and dorsolateral stripes and spots on the legs were very dark red. She has been much longer in preservative and the ground color is now a cadaverish tan instead of clear white as it is Variation. 1964a. 310. its ground color had become a pale. p. lying close together between — the moderate-sized. — When collected at night. They also occur in a narrow band along the dorsal surface of thigh. p. — The paratype is a spent. Color in pale. p. and on top of the three lateral so on sides of face toes. moderately .— Rivero. These pepper-like specks occur on dorsal surface of forearm. adult female 58. Diagnosis. 1923(a). while the frog was on a vine some six feet from the ground near a small wooded stream. The pollex. granosa from H.

skin of throat and chest smooth. reaching to disk of third which covers about two-thirds the tympanic area. was compared directly with the series of cotypes in the British BM Museum.12. femur. .6 mm. Caquetd. There seems to be no determinable significant difference in the series and we must consider all of these specimens as conspecific. roof of skull not exostosed.5 16.S. no dermal appendage on heel. the upper lip flaring out fairly strongly below it. No patagium.. in postaxillary region appreciably narrower than greatest width of head. —This tion tibia. fourth finger appreciably longer than second. a distinct oval inner but no distinct outer metatarsal tubercle. There and the only color The black . its diameter equal to its distance from nostril. 35. separated from eye by a projecting. about three-fifths the diameter of eye. no traces of a skinfold across chest. no vocal sac. when hind leg is adpressed. Eye not large but prominent.2 mm.1 individual brown above and below. arms.5. heels overlap greatly. Ecuador. hands.181 [1947. heel-to-toe. Fingers webbed at base. adult female.2. head length. when limbs are laid along the side. Color in alcohol. partially translucent upper show through the lids.. body.. which is in turn about equal the distance between nostrils. Head and body. and legs uniformly and fairly conspicuously granular. well-developed outer and inner tarsal ridges. third and fifth toes subequal. Toes slightly more than one-half webbed. Texture of the dorsal skin is somewhat reminiscent of the flesh of a stony pear. knee and elbow overlap considerably. 18. 10. palpebral membrane not reticulate. Body not elongate. Tympanum very distinct.99] from Canelos.. interorbital distance about twice the width of upper eyelid.. 24. no projecting rudiment of a pollex. disk of fourth toe covering about one-half the tympanic area.5 mm. Skin of head not co-ossified with skull. that of belly and lower surface of thigh uniformly and finely granular. eyes tend to is is is now no sign of a uniform pale grayish any pattern or pigmenta- the black of the choroid coat of the eye. the web on fourth toe reaching base of the penultimate phalanx. head — width. loreal region concave and rather oblique. 14 mm. when hind legs are bent at right angles to body. their distance eye. a well-developed ulnar ridge. third finger extending to middle of the antepenultimate pha- lanx. Remarks. mm. Dimensions. mm. hand.6 mm. —Our specimen from Florencia.12. 13. Can thus rostralis fairly well defined. heel reaches anterior margin of eye.232 U. Skin of upper parts highly glandular. NATIONAL MUSEUM BULLETIN 288 from eDd of snout about one-third that from from each other by an interval equal to about twothirds their distance from eye. the dorsal surfaces of head. feet. A rather narrow glandular ridge encircling upper part of tympanum. separated distance equal to one-half web on its own diameter. particularly with 80.

tongue two. the upper lip flaring out moderately below it.— COCHRAN FROGS OF COLOMBIA AND GOIN 233 On the other hand.16. From Hyla granosa this new species can be distinguished by the presence of dermal heel appendages and the ridge above the vent. Hyla guibei. broadly U-shaped when viewed from above. Vomerine teeth in two heavy. Narino. Eye — moderate.— CNHM 54773 from Pueblorrico-St. Colombia. collected by Gerardo Reichel-Dolmatoff. nostrils almost entirely lateral. MCZ 28049. BM 74.7. lying close together on a level with the posterior halves of the moderatesized. not particularly prominent. Para Santarem. Vaupes: Gino-Goje. their distance from end of snout about one-third that from eye. slightly rounded in profile. (cotypes). the upper jaw extending slightly beyond lower. somewhat curved. its diameter equal to its distance from nostril. Snout moderate. Lower Rio Apaporia.5-6 MCZ 2S025-37.v. broadly cordiform.10. collected in March 1962 by Gerardo Reichel-Dolmatoff.7-9 (cotypes). which is about equal distance . loreal region concave and somewhat oblique. — Diagnosis. VENEZUELA: La Culebra. Colombia. A member of the granosa group with a transverse white ridge above the vent and with triangular dermal appendages on the heels. ECUADOR: Canelos. new species Figure 22 Holotype. Specimens Examined COLOMBIA Amazon as: Cano Guacayd. separated from each other by an interval equal to about two-thirds their distance from eye. / \ series.16. Alto Cucunoma. 800 meters. Pacific side.10. east of Tumaco. in 1962 —CJG 2309. Amazonas. USNM 152756. Caldas. palpebral membrane not reticulate.thirds as wide as mouth opening. from H. Rio Satingo.12. BM 72.22. oblong choanae. Paratypes.99] (cotype).5.181 [1947.) and that the name gracilis Melin is a straight synonym of punctata Schneider. MCZ 27828. BRITISH GUIANA: Demerara Falls. Caqueta: Florencia. when the Caqueta specimen was compared with the type of Hyla granosa gracilis several differences were noted. Description of holotype. Colombia.45 (cotype). We believe that the latter is a specimen of Hyla punctata (q. Cecilia.12. BRAZIL: From interior. Narino. BM 75. its posterior border not free and unnotched. interorbital distance greater than width of upper eyelid. Rio Rosario. rubracyla it can be distinguished by the absence of dorsolateral dark stripes. CJG 2314. Canthus rostralis well defined.2. slightly projecting. BM 80.

. holotype. CNHM 54773 (same size). a low but discernible tarsal ridge. Tympanum very distinct. Skin of upper parts smooth.S. a horizontal glandular fold Figure 22. when hind leg is adpressed. web on third finger reaching base of penultimate phalanx. a short but distinct triangular dermal appendage on heel. Toes nearly completely webbed. reaching to middle of disk of third which more than covers the tympanic area. a low ulnar ridge. NATIONAL MUSEUM BULLETIN 2 88 between nostrils. separated from eye by a distance equal to one-half its own diameter. about one-half the diameter of eye. Hyla guibei. third and fifth toes subequal. projecting rudiment of a pollex slightly developed. knee and elbow overlap. when hind legs are bent at right angles to body. disk of fourth toe just snugly covering the tympanic area. when limbs are laid along the side. the web on fourth toe reaching the middle of the penultimate phalanx. heel reaches beyond tip of snout.— 234 U. fourth finger an entire disk longer than second. heels overlap considerably. very indistinct rounded outer metatarsal tubercle. No patagium. Fingers webbed at base. Body moderately elongate. in postaxillary region about two-thirds the greatest width of head. a distinct oval inner and a smaller.

The .6 mm. SQUALIROSTRIS group Members of this group are small Hyla with striped patterns. hand. and shanks having a diameter equal to that of the thighs.. We dedicate this new species to Dr. 21. 61. in recognition of our gratitude to him. a distinct narrow glandular ridge passes above skin of throat and chest. although the details of the dark spots on the dorsum are not absolutely identical. Skin of head not co-ossified with skull. This specimen is 44. head width. a distinct blotch on each upper lid that is joined to the one on the other side by an marks across the back. — Dimensions. heel-to-toe. linderi and H.. 34 mm. Variation.. head length. the most which is just above the sacrum. that of belly and lower tympanum. three dark posterior of dirty gray. femur. roof of skull not exostosed. It is impossible to tell at the present time if the differences between the female paratype and the type have any systematic significance. The three specimens at hand surely represent a distinct species in this group of white frogs of the Pacific coast. no traces of a skinfold across chest.6 millimeter in head-body length and is without dark markings except for little patches of dark brown on the heels and a dark line along the lateral margin of the fifth toe. Paris.. — 11.5 mm. tibia.2 mm. smooth. 46. surface of thigh uniformly granular. 30. Head and body.1 mm. parkeri seem to belong in this group. —A pale frog with the dorsal surfaces thickly flecked with pepper-like dots. pigmented eggs.6 mm.FROGS OF COLOMBIA — COCHRAN AND GOIN 235 passes above the vent. This female is packed with ripe. The dermal appendages on the heels are not nearly so large as those of the type. 19. Remarks. no vocal sac.. pointed snouts. the thighs otherwise being immaculate. Color in alcohol.. H. adult female. In addition to these flecks the pigment is concentrated in the following places to form a pattern: just back of each nostril a little can thai stripe that goes about halfway to eye. The pepper-like fleckings are reduced on top of the thighs to several patches. There are tiny. faint specks of pigment on the dorsal surface but these are not dense enough to keep the specimen from looking pure white to the naked eye. The female paratype is 50. Jean Guibe\ director of the Museum National d'Histoire Naturelle. 22 mm. All ventral surfaces are an immaculate interocular bar. Its pattern is like that of the type.2 millimeters in head-body length and the spiny prepollex penetrates the skin. compared —The male paratype has the ground to the cadaverish gray ground color color nearly white as of the long-preserved type. indistinct.

— MLS 122. Lutz Figure 23 1925. 1963.— Gorham. p. and we NATIONAL MUSEUM BULLETIN 288 Only one species refer it to squalirostris group but is is here kept separate now known from Colombia. Hyla squalirostris is perhaps more closely related to H. Vomerine teeth in two small rounded series. Lutz. . is closely related to the rubra for convenience. Cochran. MLS 122 (X 2. boundary of Rio de Janeiro and Sao Paulo) 1926. 23. 151. squalirostris A. 212 (type locality. 14. Norte de Santander. 1955.— 236 group U. Hyla squalirostris A. A small. p.— Hyla squalirostris. but its slender build and lack of black and light markings on the rear surface of thigh serve to distinguish it from that species. Description. marshes of Serra da Bocaina.S. rubra than to any other Colombia frog.5). from 20 kilometers west of Cucuta. Colombia. Hyla . p. pending study of additional material. Figure 23. slender frog with long legs and a long pointed snout and longitudinal stripes on the back. — Diagnosis. p. 7.

no project- ing rudiment of a poll ex. separated from each other by an interval equal to about two-thirds their distance from eye.5 mm. 4. cordiits posterior border very slightly free and very shallowly notched.. separated from eye by a distance equal to one-fourth its own diameter. a pair of lateral stripes originate on the tip of the snout. tibia. interorbital distance greater than width of upper eyelid. Tympanum distinct. its diameter equal to its distance from nostril. no dermal appendage on heel. fourth finger but slightly longer than form. when limbs are laid along the side. 5. their distance from end of snout about one-third that from eye. adult male. considerably projecting. head width.5 mm. a distinct oval inner and a much smaller but distinct rounded outer metatarsal tubercle. striped frog. each passes posteriorly through the nostril. 12. 6.. 15. Dimensions. tongue one-half as wide as mouth opening. that of belly and lower surface of thigh uniformly and finely granular. no ulnar ridge. Fingers without webs. heel reaches tip of snout. knee and elbow overlap considerably. heels overlap slightly. second.6 mm. third and fifth toes subequal. Snout long. traces of a skinfold across chest. rounded choanae. through brown with five longitudinal . skin of throat and chest smooth. 8 mm.. vocal sac apparent as subgular wrinkles.FROGS OF COLOMBIA —COCHRAN AND GOIN 237 lying close together on a level with the posterior borders of the small. pointed in profile.8 mm. when hind leg is adpressed. Head and body. loreal region flat and very nearly vertical. in postaxillary region a little narrower than greatest width of head. Body elongate. — Color in alcohol.9 mm. failing to reach to disk of third by the diameter of a disk. the upper jaw extending considerably beyond lower. the web on fourth toe reaching the middle of the antepenultimate phalanx (webs very delicate and tear at a touch). a rather narrow glandular ridge encircling upper part of tympanum. No patagium. disk of fourth toe covering about one-fifth the tympanic area. 9. The dorsum is medium dark stripes as follows: a medium stripe originates between the eyes and terminates posteriorly above the urostyle. Toes about one-half webbed. the upper lip not flaring out appreciably below it.. —A dark. pointed when viewed from above. when hind legs are bent at right angles to body. disk of third covers about one-fourth the tympanic area. heel-to-toe. roof of skull not exostosed. Skin of head not co-ossified with skull.1 mm. palpebral membrane not reticulate. femur. Eye moderate in size but prominent.. hand. f along the canthus rostralis. Skin of upper parts smooth. a pair of dorsolateral stripes originate one behind each eye and terminate posteriorly to the sacral hump on each side. head length. no tarsal ridge. Canthus rostralis well defined.. nostrils more lateral than superior. which is relatively narrow and slightly less than distance between nostrils. about three-fifths the diameter of eye.

— Specimens Examined COLOMBIA 20 km. if not identical with. Serra da Bocaina (boundary of Rio de Janeiro and Sao Paulo). We have tried to of these geographic variations as simply populations of rubra and have restricted nomenclatorial recognition to distinct. vomerine teeth in two small rounded patches. sensu latu. our specimen is certainly close to. or tarsal ridges. west of Cucuta. rubra. as is the side and venter below the lateral stripe. There seems to be no doubt that squalirostris is allied to parkeri Gaige. to the rubra group.S. However. the Brazilian specimens of the species. for the Serra da Bocaina and the Maracaibo Basin have little in common faunistically. Bonita. The single specimen at hand seems to differ structurally from the Brazilian specimens only in that the snout is a little shorter and the tympanum a little larger and set a little closer to the eye. The three species now known from Colombia may be distinguished by the following key: . than squalirostris and parkeri are to either of them. and lindneri Miiller and Hellmich and. however. NATIONAL MUSEUM BULLETIN 2 88 lower part of eye.238 U. we do not feel that this justifies putting these species in the rubra group as at present understood. sympatric populations that show an obvious relation to rubra. Maschwitz. Norte de Santander: ARGENTINA: BRAZIL: USNM USNM 96721. 96719-20 (cotypes of squalirostris). for it seems probable that such forms as rubra and hayi are much more closely related. Buenos Aires. rubra group this is one of the most difficult groups among the South American tree frogs. MLS 122. evelynae Schmidt. 96608-13. is one of the most widespread and one of the most variable of all South American frogs. through them. 12. ulnar. Sao Paulo. The edge of the upper lip is pale. The group as a whole is represented by small to medium-sized Hyla with reduced webbing between the fingers. It bothers our zoogeographic sense somewhat to use the name "squalirostris" for this single specimen. and by the presence of distinct markings on the posterior surfaces of the thighs. as they are based on comparison of a single specimen with a population about which we know little concerning geographic variation. one to the other. H. in consider all some cases quite markedly. the absence of patagia. Every local population differs in some respects from other At the present time all local populations. It is impossible to judge the significance of these differences. Remarks. MCZ 35421-2. and through the tympanum and terminates in the groin.

1935.— 45. p. Pebas. rubra Hyla rubra Daudin Plate 31d-f 1802. 1862. 1870. 1939. 526. locality. . p. p. 413. 1830. locality.— Lutz. p. 124. rubra. p. 51. Cope. and rostrum pale H. 1959. — Reinhart and 1882a. 1820. 310. p. p. Underside of throat. 66. 109. 1830. 1964b. Boettger. 1877. 1937. p. 1838. 35. p. p. 1937.— Barbour and Loveridge. 266. p. Hyla 1951. p. p. 33 (type locality. 110.— Estable. chin. 18. p. 1873a. 558 (type locality. 1830.— Gunther. Nauta). vertical bands. Scytopsis cryptanthus Cope. 448. 1874b. p. ex Ecga von Solimoens). 108 Fowler.— Peters.— Beebe.— Fitzinger.— Cei. 89. p. p. 1888a. 1830. 1958. p. 1860. 91 Parker. 1892. 403. 464 (type locality. — p. p.— Berg. 61. (type locality. 60.— Mello-Leitao. foliamorta Markings on posterior surfaces of thighs in the form of vermiculations. — Peters. Scytopsis ruber.— Steindachner. — Wagler.— . p. Bahia). 16. p. 1858. Tschudi. p. 226. 1926. p.— Nieden. 1912. 35. 705. 13. 1874. and Loveridge. p. p. 214. 123 (type locality. 1871a. p. 1913.— Frieberg.— Crawford. — Gunther. a 6 l b 2 . — — 1874b. 1873a.— Gorham. p. p. 1841. Rivero. 171.— De Witte. 156 45— Barbour 1874.— Baumann. p. 122. 311. . 638. — Schmidt and Inger. 1873a. 1963. Liitken. p. 1803. 1858. 1933. — COCHRAN of thighs in the AND COIN form 239 of broad. p. 1874b. 120. p. p. Pard. 1904. 1958. "Guyane")- 1863. 162 (type locality. 1820. 123.— Travassas and p. p. p. p. 23. p. 1933. 1160 Miranda-Ribeiro. Hyla x-signata Spix. 1919. 1939. p. 1863b. p. 1959b. p. conirostris 613. Burmeister. 1856. — — 1956. 1942. 208. 201. 1928. Peters. 222. 1929. 1838. 299. Muller.— Gorham. 172. p. H.— Goin. 1884a. 201.— Mertens.— Goin and Layne. 602. — Cope. 1935. 1874. — Andersson. 151. 218. Merrem. p. p. 1961. Werner. p. p. — Peters. 207. 389. p. 1929. p. Hyla afinis Spix. 1930. Stebbins and Hendrickson.— Carvalho. 53. 1871b. p. Suriname). dark. Auletris coerulea. p. p. 1913.— Cott. Scytopis alleni Cope. p. Hyla coerulea Spix. — Peters. 104.— Liu. 37 (type locality. Hyla 1870. — Boulenger.— Lutz. p.— Crawford and Jones. 1901. Auletris 40 (type p. Scytopsis rubra x-signatus. 1949. 1942. p. Calamita ruber. Ecuador). 23. 110. boulengeri Underside of throat. p. Dendrohyas rubra. 1927.— Dunn. 338. Hyla quinquefasciata Fowler. 1963. 275. 160 (t}'pe Nieden. 19 p. 4. p. 413. p. 1919. 444. 174 Ruthven. p. 124. 1952. 268. 1824. 403. 1886a. 50. 56. Hyla cynocephala Dunieril and Bibron. 1867. 280. p. 39.— Lidth. p. 32 (type locality not given). 1896. Peters. 40. p. p. 2 . and rostrum dark H. 1884b. 1873a.— Wagler. p. . 156. p. p. p. — 1824. p. p. p. Peru). 340. p. p. 1951b. 1927. 512. Hyla rubra Daudin. 1931. 1942. 1931. 196. p. p. 1873a. p. 120. 1874b. p.— Goin. 1926. 1923.— Schubart.— Guib<§.— FROGS OF COLOMBIA a Markings on posterior surfaces l . Peters. p. 282. p. p. p. 1923. Amazonenfluss) — Peters. p. — — — — Frietas. p. chin. 74. 1945. Chimbo. 94. 280. 218. 1939. 1824. Brazil) (nomen nudum). p. 1892. p. 88. lateristriga Spix. 1925. 1824.

Hyla rubra duartei B. no projecting rudiment of a pollex. 32 (type locality. rubra likely to —CNHM . Colombia. a distinct oval inner and a much smaller but distinct rounded outer metatarsal tubercle. when hind legs are bent at right angles to body. prominent. 34 (type locality. separated from each other by an interval nearly equal their distance from eye. NATIONAL MUSEUM BULLETIN 288 Hyla rubra hubneri Melin. third and fifth toes subequal. broadly rounded. somewhat pointed when viewed from above. 301. no ulnar ridge. Eye moderate. short-legged Hyla with just a trace at base of fingers on posterior face The and with distinct black and white markings of thighs. rounded series. 1951b. are H. 1951. no tarsal ridge. the web on fourth toe reaching nearly to base of the penultimate phalanx. Canthus rostralis slightly defined. p. knee and elbow fail to meet.800 hi. which closest relative of it is most H. lying close together between the small. 1951. 1960. boulengeri and H. p. Aniazonas. 1941. Body not elongate. Description. from Mariquita. Hyla rubra nasica. in postaxillary region a little broader than greatest width of head. when limbs are laid along the side. Taracua. interorbital distance about equal to width of upper eyelid. fourth finger considerably longer than second. broadly pointed in profile. 81818. Alto Itatiatia. Brazil. its diameter equal to its distance from nostril. U. heel reaches anterior margin of eye. disk of fourth toe covering about one-third the tympanic area.). their distance from end of snout about one-half that from eye. just reaching to disk of third which covers about one-half the tympanic area. Fingers with a trace of web at base. the upper lip flaring out below it. flat. the upper jaw extending considerably beyond lower. — Diagnosis. its posterior border slightly free and almost imperceptibly notched. joliamorta. Hyla rubra inconspicua Melin. 705 (type locality. which is relatively wide and slightly greater than distance between nostrils. very slightly projecting. of web —A moderate-sized. Tolima. nostrils between lateral and superior. Rfo Vaup6s. Lutz. no dermal appendage on heel.S. vertical dark bands. when hind leg is adpressed. about one-half the diameter of eye. Toes slightly more than one-half webbed. separated from eye by a distance equal to one-third its own diameter. rounded choanae. From both of these it may be distinguished by its smaller size and the fact that the black markings on the face of the thigh are in the form of vermiculations rather than broad. Peru). p. Tympanum very distinct. above 1. tongue three-fourths as wide as mouth opening. Hyla rubra fuscomarignata B. loreal region rounded and somewhat oblique. p. in Colombia. 1960. Lutz.240 1941. Brazil). Vomerine teeth in two short. palpebral membrane not reticulate. and the ones with be confused. p. Roque. Snout moderate. Gans. 706. 1951b.

Head and body. Dunn told us many years ago that when he first started studying the lizards in the Institute deCienciasNaturales.6 mm. 11 mm. a line from the posterior corner of the eye that passes above the tympanum.2 mm. in this species.. The area on each side between the dark lines that terminate above the shoulder and those that pass on back to the sacral hump is somewhat lighter than the rest of the dorsum and appears to form an indistinct dorsolateral light stripe. 10. the — rest of the ventral surface — is a dirty gray. as we have done above. Each groin and the anterior face of each thigh is marked with distinct black flecks and the posterior surface of each thigh with distinct. he would take a bottle of one species off the shelf and find that it had been described six times and then find that the next six species had not been described at all. 16. it seems to us rather futile to attempt to apply names to each of the populations. in Bogota.. a well-developed skinfold across chest. There are a few faint fleckings on the chin and throat. In the case of the present species.. as we now understand it. While of course the number of times mentioned by Dr. heel-to-toe. 12 mm.. somewhat vermiculate areas of light that are outlined with dark brown and which form the thigh pattern so characteristic of this species.7 mm. 13.5 mm. Skin of upper parts smooth. widespread species of South American amphibians and reptiles described time and time again.. 1958) and that each of the populations . A rather brownish frog with indistinct dorsolateral stripes.9 mm. we do not mean to imply that rubra is not a variable species with perhaps a number of geographic races worthy of recognition. roof of skull — head not not exostosed. tibia. No —COCHRAN AND GOIN 241 patagium. an inverted triangular-shaped interocular bar. By placing all of these names in the synonymy. Dunn in his story is fictitious. both geographic and individual. Remarks. thence downward toward the shoulder where it terminates. we can state that the specimens from Leticia have a different tooth count from those of Trinidad (Goin. On the moderate brown dorsal background indistinct darker brown markings occur as follows: a narrow line along each canthus from naris to eye. Skin of with Dimensions. femur. head length. vocal sac subgular. for we find common. From our experience. head width. skin of throat and chest very finely granular. adult male. 23. Color in alcohol. the implied situation is not. we find that a new name has been proposed for it (or a part of it) no less than 14 times. a line from the hind margin of the upper lid that passes straight back to the region of the sacral hump.. hand.FROGS OF COLOMBIA heels overlap greatly. However. co-ossified skull. a rather narrow glandular ridge encircling upper part of tympanum. that of belly and lower surface of thigh uniformly and not so finely granular. until a really thorough study is made of the variations. 32.

Santander: 4. cynocephala. 8516. from which we have seen fresh or Moscopan live material —Amazon basin. 152196-7. USNM 152123-7. 8511 (3). Beltran. CJG 2351-2. 146258. MLS 197. Rfo Meta: Caiio Losada. CNHM 61753. dencia. 2330 (40). Florencia.71. and. 20 km. "Colombia": BM 1915.10. east of Bogota.21. We and and quinquefasciata. Putumayo: Puerto Asfs. CNHM 5647. MLS 28. 29a. MLS 29. Serrania de La Macarena. Rfo Orteguaza. Cundinamarca: Aguadita. 82.S. 21. Rio Vaupes. the rainforest in the lowlands of western Colombia. 152191-2. Sincelejo-Tolu. CJG 2458-9. JRT 394-7. Morelia.71. was correct in using We have studied a cotype 301) as a race of this species. CNHM 81818-20. and Bolivia has a somewhat different mien. Moscopan. C6rdoba: Rio Manso. MLS 73. MLS 280. BRAZIL: Para. CNHM 54777. MLS 14. Tierra Alta. just as obviously. Puerto Narino. 8512 (2). Rio Zulia. CNHM 54774. UF 8510 (3). Andes. USNM 152282. region of Guamo. there is much type material that needs to be studied. 476 (41-7893) (type of Hyla rubra hubneri). 152000-9. west of Cucuta. USNM 151452. 2272 (10). inconspicua. NM .7 km. do not think that Gans (1960. ANSP 2159 and MCZ 1519 (types of Scytopis alleni) Jaracua. CJG 2432 (113). USNM 152120.d. 25472152075-80.10.21. Caqueta: Aserrfo. Paramo de Tama. Amazonas: Boyaca: Muzo. Rio Rumiyaco. southeast of Bucaramanga. ANSP 23343-5. USNM USNM USNM USNM 152247. Villavicencio. USNM USNM Cauca: Choc6: Andagoya. ANSP 24398-400. . Norte de Santander: La Selva. USNM 152072-4. Guayabero. Mesa de Cucuana. CJG 2329. CJG 2269-71. Valle: Rfo Rapaso. 5835) from Paraguay and do not believe it is of the species rubra. USNM 146255. Near Altimira. Serrania de La Macarena. Caldas: Pueblorrico. hubneri. One of the reasons we are not ready to commit ourselves to the assignment of names to the different populations is because obviously there are many large and important geographic areas from which we have not seen well-preserved fresh material. Narino: Imbili. La Provi152038-46. USNM 152312. 8515 (3). Suriname. 8513 (11). Htjila: Neiva. Specimens Examined COLOMBIA Leticia. 152288-311. We have seen the types of only five of the named forms listed in the synonymy: Trinidad. MLS 212. CJG 2443. Rfo Orteguaza. species. in the — alleni. ANSP 24332-4.242 NATIONAL MUSEUM BULLETIN 288 U. BM 1915.c. We Cope's name nasica of nasica (USNM p. therefore leave the problem of working out the various races sibling species (if any) of rubra to someone who has the time and material to tackle the whole problem throughout the range of the We hope that this will be soon. FM 16-21. BoLfvAR: Gambote. 28a-c. Mariquita. Amazonas. Tolima: Espinal. 152184. 8514 (11). Villa Marfa.

31 (type locality.— Nieden. 339. Perlata. FROGS OF COLOMBIA — COCHRAN . 1901. p. considerably projecting. 37863. p.— Dunn. — A rather large Hyla of the rubra group with broad. Hyla boulengeri. vertical dark bands rather than in the form of vermiculations. Hyla boulengeri (Cope) Plate 32d-f 1887. dark bars on the posterior faces of thighs and generally with Diagnosis. CJG 2391-4. interorbital greater than width of upper eyelid. El Real. NM UF TRINIDAD: 3648 (4).— Rivero. 279. separated from eye .— Gaige. 64003. "GUIANA": MHNP 765 18115 (type of Hyla quinquefasciata) (type of Hyla cynocephala) USNM USNM PANAMA: 140569-70. 413. Trinidad River. very elongate choanae. p. Costa Rica). Hartweg. Hyla lancasteri Barbour. the upper lip flaring out strongly below it.. Panama Canal Zone. From its close relatives in Colombia H. 311. p. 1931. not particularly prominent. separated from each other by an interval equal to about two-fifths their distance from eye. 1964b. SURINAME: Paramaribo. 1923. rubra by having the dark markings on the posterior surfaces of the thighs in the form of broad.— Gorham. the upper jaw extending somewhat beyond lower. palpebral membrane not reticulate. 1958. Colombia. 1963. p. rounded in profile. 199. its posterior border fused and with just a trace of a notch. Nicaragua). For a detailed comparison of boulengeri with joliamorta see "Remarks" under the latter species. 125. tongue two-thirds as wide as mouth opening. p. and the throat pale rather than dark. 1918. 1928. lying close together between the moderate-sized. p.— Noble. 21. Description. 1960a. their distance from end of snout about one-fourth that from eye. loreal region concave and rather oblique. 12 (type locality. AND GOIN 243 ANSP ECUADOR: Chimbo. p. Scy topis boulengeri Cope. Eye moderate. p.— Barbour and Loveridge. Tympanum is distance slightly in turn equal to the very distinct. broadly rounded. 1887b. Barbour and Loveridge. nostrils more lateral than superior. 856 Fouquette. Darien. 1929. USNM — . Snout elongate. 1937.— Giinther. 480 (type of Hyla rubra inconspicua) PERU: Roque. p. 1952. boulengeri may be distinguished from H. which distance between nostrils. Choco.— Taylor. vertical a rather light chin and throat. from Andagoya. 267. p. 1929. two transverse series. and from H. p. — 1963b. p. its diameter equal to about three-fifths its distance from nostril. 140573. 251. Canthus rostralis slightly defined. 4. about three-fifths by a distance equal to one- the diameter of eye. rather lanciform when viewed from above. 279. p. p. —CNHM Vomerine teeth in 81854. joliamorta by having the undersurface of the proboscis. and Stuart. 1901. the chin. 58.

no tarsal ridge. third and fifth toes subequal. no dermal appendage on heel.7 millimeters in head-and-body length. head width. 14. Fingers with just a hint of 2 88 web at base.. in postaxillary region a little narrower than greatest width of head. a transverse bar at the level of the sacral hump. knee and elbow barely overlap. and two little longitudinal bars between the sacral hump and the vent. — . 12. no ulnar ridge. a distinct oval inner and a smaller rounded outer metatarsal tubercle. head length. The type has faded to the point where no pattern is discernible but it is perfectly recognizable structurally. Body moderately elongate. and then fades out just posterior to the suprascapular region.2 mm. that of belly and lower surface of thigh uniformly and finely granular. 36.5 mm. and these bands continue — — on to the posterior surfaces of the thighs. The amount and degree of variation in the specimens before us are hardly more than would be expected in any lot of specimens collected and preserved by different persons at different times. hand... vocal sac not expanded. the web on fourth toe reaching the distal end of the antepenultimate phalanx. adult male. a band on each side that originates behind the eye. when limbs when hind are laid legs are bent at right angles to body. when hind leg is adpressed. Color in alcohol. The specimens from Colombia range from 32.. 21. 27. not reaching to disk of third which covers about one-third the tympanic area. along the side. own NATIONAL MUSEUM BULLETIN diameter. heel reaches tip of snout. fourth finger just slightly longer than second. passes above the tympanum. the ventral surfaces are immaculate. femur. A medium brown frog with mottlings of dark brown. Skin of head not co-ossified with skull. Toes slightly more than one-half webbed.6 mm. tibia. Except for a faint hint of dusky pigment around the edge of the lower jaw. 17. roof of skull not exostosed. The most conspicuous of the darker brown dorsal markings are as follows: an interocular triangle with its apex directed posteriorly.5 mm. heels overlap greatly. a rather narrow inconspicuous glandular ridge encircling upper part of tympanum.2 mm. heel-to-toe. no projecting rudiment of a pollex. Skin of upper parts very finely granular on top of head and snout and in the region behind the tympanum and above the shoulders but otherwise the dorsal surfaces are rather smooth.244 half its U.2 mm. skin of throat and chest smooth... Most of the rest of the pattern is simply made up of tiny flecks of pigment scattered over the medium-brown ground color. Remarks. Head and body. 10. Dimensions.5 mm. well-developed traces of a skinfold across chest. a small L-shaped blotch in the middle of the back. disk of fourth toe covering about one-third the tympanic area. No patagium.S. There are several broad dark bands across the thighs and shanks.9 to 48.

Hyla foliamorta Fouquette. foliomarta can be separated from H. boulengeri. Description. it may be differentiated by the dark throat and chin and the dark underside of the rostrum (see "Remarks" for a detailed comparison). 70a. Antioquia. MLS 211. pointed when viewed from above. Norte de Santander: La Selva. MLS 118. slightly V-shaped series. Snout long. from El Real. of Miraflores Locks. USNM 152285. A detailed comparison between boulengeri and foliamorta under the discussion of the latter species. loreal region not concave but very oblique. the upper jaw extending considerably beyond lower. p. northwest Zone). its posterior border very slightly free and unnotched. vertical bars rather than vermiculations. nostrils more lateral than superior. dark bars on the posterior face of the thigh and with generally a dark chin and throat and with the ventral surface of the rostrum Diagnosis. CNHM 81854-5. their distance from end of snout about one. lying close together between the moderate-sized. Rio Piraboton. — USNM 127865. tongue twothirds as wide as mouth opening. is given Specimens Examined COLOMBIA Choc6: Andagoya. oblong choanae. Maraquita. Cttndinamarca: Caparappf. vertical rather dark. Vomerine teeth in two short. 125 (type locality.FROGS OF COLOMBIA The specimens from to be different in any the —COCHRAN AND GOIN 245 drainage in Meta do not seem manner from the other Colombian Amazon significant specimens. H. C6rdoba: Rio Manso. seven mi. USNM 152186-8. Putumayo: Puerto Asfs. Villavicencio. the upper lip 837-262—70 17 . Panama Canal — A rather large frog of the rubra group with broad. to which it is very closely related. 82008. rubra by having the dark markings on the posterior faces of the thighs in the form of broad. pointed in profile. Colombia. USNM 152278. broadly rounded. Can thus rostralis slightly defined. Santander: Simitarra. 13974 (type). From H.third that from eye. separated from each other by an interval equal to about one-half their distance from eye. considerably projecting. Mouth Meta: of Cailo Losada. CNHM CNHM 81824-5. CJG 2431. MLS 70. NICARAGUA: Tolima: Vaupes: USNM Hyla foliamorta Fouquette Plate 32a-c 1958. Rio Guayabito.

about three-fifths the diameter of eye. adult male.. joliamona was made by the junior author and Dr. Tympanum very distinct.. no dermal appendage on heel. no ulnar ridge. heel-to-toe.7 mm. . The entire undersurface of chin and throat is tinged with dusky. NATIONAL MUSEUM BULLETIN 288 below it. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. 26. heels overlap greatly. J. No patagium. 14. no tarsal ridge. that of belly and lower surface of thigh uniformly granular. in general although not invariably. Toes slightly more than one-half webbed. 34. A dark brown frog with a limited pattern of Color in alcohol.. Dimensions. when limbs are laid along the side. the web on fourth toe reaching the distal end of the antepenultimate phalanx. separated flaring out rather strongly little less from eye by a distance equal to one-half its own diameter. not reaching to disk of third which covers about three-fourths the tympanic area.. roof of skull not exostosed. "Comparison of joliamorta with boulengeri: First. when hind legs are bent at right angles to body. This can be more readily seen from the ventral view. 40. in postaxillary region a little narrower than greatest width of head. no projecting rudiment of a pollex. palpebral its diameter a membrane not reticulate.6 mm. femur. on the wrists there are faint crossbands of grayish brown. hand. head length. Head and body. — while they had all of the Colombian material of boulengeri and foliamorta and the type oi Joliamorta before them. a skinfold across chest. black supratympanic glandular ridges. to a lesser extent.3 mm..3 mm. Skin of head not co-ossified with skull. head width. The following comparison of Hyla boulengeri and H. third and fifth toes subequal. fourth finger but slightly longer than second. a pair of small but conspicuous suprascapular spots.S. and at insertion of the arms. knee and elbow overlap considerably. than its distance from nostril. M. Fingers with but the merest trace of web at base. A well-developed narrow glandular ridge passes above the tympanum skin of throat and chest smooth. Body moderately elongate. Remarks. 15.3 mm. tibia. 21. Fouquette black. no vocal sac apparent. above the tympanum. disk of fourth toe covering about twothirds the tympanic area. Eye neither large nor prominent. Skin of upper parts generally smooth but with a few granulations on top of snout.7 mm.246 U. — — The main elements of the pattern are as follows: a blackbordered interocular triangle with its apex pointed posteriorly.. when hind leg is adpressed. in the region behind the eyes.8 mm. On top of the thighs and shanks and. joliamorta seems to have more pointed tip to the snout. and black bands across the anterior and posterior faces of the thighs. 12. . interorbital distance slightly greater than width of upper eyelid and than the distance between nostrils. heel reaches beyond tip of snout.

as do the three suprascapular spots posterior to it. were the only known Colombian representatives of foliamorta. the corner of the groin in boulengeri is is more more often dark. more tuberculate skin. The dark interocular triangle in foliamorta seems to be bordered by a narrow but quite distinct glandular ridge on all three sides. whereas in foliamorta the lower margin of the lower jaw seems to be rather smooth all the way from front to back. These same stripes and spots can usually be made out in boulengeri but because of the dark roughened skin they are not nearly so prominent. the leg of boulengeri seems to be shorter. An additional character can sometimes be noted along the lower jaw. USNM 127865-6. the dusky spots are so small and so thickly scattered that you get the impression of a more or less gray or dark throat. the heel reaching nostril. the corner of the groin in foliamorta often clear. it seems unlikely that their present distribution can be worked out in any detail except by studies made on the species in life. Antioquia. the dark supratympanic stripe that follows along it seems to stand out rather strikingly against a generally paler background. whereas in boulengeri there tends to be in this region a dark spot similar to those along the face of the thigh. brownish perhaps.FROGS OF COLOMBIA —COCHRAN AND GOIN 247 It is a more projecting snout. whereas in foliamorta it tends to be longer. boulengeri seems to have a rougher. the heel reaching to the nostril or to the tip of the snout or perhaps even somewhat beyond that. Also. In foliamorta. between the eye and in boulengeri but when they are present they are never so conspicuous. Colombia. In other words. ." At that time it was decided (by Goin and Fouquette) that the two specimens listed here from El Heal. Dorsally. If these two nominal forms are really distinct. This ridge is not actually in contact with the dark triangle but separated from it by a millimeter or two. Secondly. Furthermore. In general. whereas foliamorta seems to Sometimes similar ridges are present have the skin of the dorsum smoother and less tuberculate. on the postocular glandular fold of foliamorta. males of foliamorta seem to have a darker throat than do males of boulengeri. From a ventral view the lower surface of the rostrum tends to be dark in foliamorta whereas generally it is light with perhaps just some dark flecks on it in boulengeri. Specimens Examined COLOMBIA Antioquia: El Real. Sometimes in boulengeri these stripes and spots may not even be discernible. whereas in boulengeri you get the impression of a white throat with dark specks scattered about on it. in boulengeri this is usually margined with tubercles ranging from inconspicuous to conspicuous. foliamorta tends to have the region of groin where the anterior face of the thigh meets the side clear.

24 (type locality. Peru). —USNM 152757.— Gorham. subocular blotch runs from the eye to the lip on each side. and with distinct crossbands on the thighs. — 1963. with distinct dorsal patterns. 1963. Hyla mimetica Melin. short. Hyla palpebrogranulata Andersson. Description. 22. The form and position of the vomerine teeth somewhat resemble those in Cryptobatrachus and Hyla phantasmagoria. the upper jaw extending a little beyond lower. PALPEBROGRANULATA group group are moderate-sized Hyla with the vomerine teeth in two. —A moderate-sized Hyla with the vomerine teeth in two with the posterior margin of the choanae).248 U. palpebral mem- . 22. U-shaped when viewed from above. There is a distinct. p. distinct crossbands on thighs and shanks. The group is presently represented by only one species. broadly rounded. its diameter not quite equal to its distance from nostril. A distinct. white. Roque. Cundinamarca. Gorham. p. transverse series. p. hardly at all superior. Eye neither large nor prominent. 14 (type locality. Snout moderate. short. lying close together on a level with the posterior borders of the small. Colombia. transverse series (on a level leucophyllata group. tongue two-thirds as wide as mouth opening. throat. and venter. the upper lip flaring out slightly below it. with well-developed patagia but without ulnar or tarsal ridges or dermal heel appendages. from Medina. somewhat projecting. Chaco. but palpebrogranulata does not seem to be closely related to either and can be immediately distinguished by its patagium. p. 13. their distance from end of snout about onethird that from eye. Canthus rostralis moderately defined. 1941.S. its posterior border fused and unnotched. transverse series on a level with the posterior margin of the choanae. rounded choanae. NATIONAL MUSEUM BULLETIN PANAMA CANAL ZONE: 23109 (type of Seven mi. reduced webbing between the toes. Members of this Hyla palpebrogranulata Andersson Plate 33a-c 1906. a well-developed patagium. Vomerine teeth in two short. The patterned thighs and the granulations on the back separate it at once from the members of the Diagnosis. separated from each other by an interval equal to about two-thirds their distance from eye. and dark flecks on chin. Bolivia). northwest Hyla foliamorta) 2 88 of Miraflores Locks. nostrils lateral. slightly rounded in profile. Tatarenda. TNHC . white. subocular blotch on each side that extends from eye to the margin of the Up. loreal region slightly concave and a little oblique.

no distinct traces of a skinfold across chest. rounded granular warts. Fingers webbed at base. separated from eye by a distance equal to one-half its own diameter. heel-to-toe.FROGS OF COLOMBIA —COCHRAN AND GOIN 249 brane not reticulate. no projecting rudiment of a pollex. dark crossbands occur on the dorsal surfaces of shanks and less-well-defined but nonetheless distinct ones occur on dorsal surfaces of thighs. A brown frog with dorsal markings of darker brown. The ventral surfaces are a dull brownish gray. A small patagium extends from the back of the upper arm to the side of body.3 mm. hand. 17. no tarsal ridge. disk of fourth toe covering about one-half the tympanic area. The posterior and anterior faces of thighs are without pattern. Broad. web reaching to middle of antepenultimate phalanx on lateral side and middle of antepenultimate phalanx on medial side. head length.4 mm.2 mm. 36. 14. tibia. knee nearly reaches axilla. skin of throat and chest fairly smooth. but more widely scattered ones occur on top of snout and shanks. no ulnar ridge. Body not elongate. disk of third finger very nearly covers the tympanic area. Top of snout and loreal regions are somewhat darker than the rest of the dorsal surfaces exclusive of the blotches. Skin of head not coossified with skull. 12. — — . that of belly and lower surface of thigh uniformly granular.5 mm. Head and body. fourth finger a disk shorter than third but a disk longer than second.. about two-thirds the diameter of eye. 11. head width. A dark interocular bar connects posteriorly with a dorsal blotch that extends backwards to the vicinity of the sacral hump. heels overlap greatly. which is in turn about equal the distance between nostrils. femur. A dark postocular bar extends from eye through the tympanum and fades out above the shoulders. roof of skull not exostosed. Color in alcohol. 25. Skin of upper parts with numerous small.7 mm. a distinct oval inner and a smaller but very distinct rounded outer metatarsal tubercle. vocal sac slightly discernible at angle of jaws. heel reaches tip of snout.8 mm. interorbital distance slightly greater than width of upper eyelid. no dermal appendage on heel. a well-developed glandular ridge passes above the tympanum. Tympanum very distinct. when limbs are laid along the side.. There are a few small blotches on the back posterior to the sacral hump and a few small rounded blotches high up on the sides between the axilla and groin. adult male.. third and fifth toes subequal.3 mm. when hind leg is adpressed. the web on fourth toe reaching the middle of the penultimate phalanx.. Dimensions. 20. These warts are most thickly concentrated on top of head and dorsum. in postaxillary region a little narrower than greatest width of head... when hind legs are bent at right angles to body. well-developed. A light subocular area is reminiscent of that of Hyla versicolor. Toes slightly more than three-fourths webbed.

S. in Medina. the same granulated skin on the dorsum. the interocular bar. and it seems to us that there can be no doubt that the two specimens are conspecific. the same vomerine tooth pattern. 38. and essentially the same amount of webbing between the fingers and toes. Both types and we believe it represents the . but we can see no other structural difference between palpebrogranulata and mimetica. from Peru. The patterns of the two specimens are not identical but there is no more difference between them than there — is between any of those in a series of six specimens (USNM 152092-7) taken at the same time and same place. to USNM 152091. these being much more distinct on the shank than on the thigh. making the dorsal surface very distinct some specimens granular. In others the ventral neckings are reduced. particularly posteriorly. rounded. The specimens range from such extremes as USNM 152090. The amount of the ventral markings is quite variable. brown blotches on the back. Both that type and USNM 152091 seem to have the dorsum smooth rather than coarsely granulated as in palpebrogranulata.5. reminiscent of the condition seen in of Osteocephalus. NATIONAL MUSEUM BULLETIN 288 — All the specimens we have seen have dark blotches on but no two have exactly the same pattern. Remarks. in which there is a solid. In some individuals there are distinct dark spots along the lower margins of the jaws. Our three largest specimens are 36. which leaves the areas lateral to the dorsal dark blotch paler and thus reminiscent of dorsolateral stripes. The dorsal body surface in most individuals is thickly covered with tiny. the dorsal pattern is quite variable.5. The specimen described above has been directly compared with the type of palpebrogranulata in the Zoological Institute of the University of Stockholm. which has scattered. the tympanum is dark. otherwise. We have also examined the type of Hyla mimetica Melin. The dorsal crossed with about three rather and rather broad dark bands. Although the described specimen is a little larger and a little darker than the type. In every specimen there is a distinct white blotch on the upper lip below the eye. these differences seem to be due to age and preservation. which is now preserved in the Zoological Museum of Uppsala. They have the same general build. and 40. wart-like structures. In every individback. and anterior chest region. and the postocular region around dark is a ual there Variation. dorsal dark blotch from between the eyes to above the sacral hump. median.250 U. same species.3 millimeters in head-body length. Cundinamarca. but in nearly every individual some degree of ventral surface of each thigh and shank necking is is discernible. but nonetheless the thigh is heavily colored rather than clear. small. the throat region.

72. The length of the crus not more than half the head-body length. 132. p. H.— Stebbins and Hendrickson. p. 8. 77. —Hellmich. 522. 261. 1944a. 40. p. p. 263. 131. mountainous surroundings 618. p. H. 573. Medium-sized Hyla with the vomerine teeth in two small.— Rivero.— Nieden. 10. Hyla . 132. 1916. labialis krausi Posterior face of the thighs distinctly mottled . — Stebbins and Henderickson. p. 10. Cundinamarca: Medina. 397. p. Hyla labialis Peters. 1959.— Stebbins and Hendrickson. — Peracca.—— .— Dunn. labialis.— Dunn. p. 1959. 1899. p. p. Nieden. 251 each upper of lip. 1. 1959. ZIMUG 41/7893. rounded patches and with well-developed patagia which are bright blue in life. p. p. p. 1923. FROGS OF COLOMBIA — COCHRAN AND GOrN have the distinct white blotches on the margin between the edge of the lip and the eye. of Bogota). p. Bogotii). p. 1963a. — Lutz.— Nieden. H. 263. 55) reported from Santa Marta. USNM 152054-5. 1923. — Dunn. 1961. 152757. 1873b. 1927. 1940a.— Stebbins and Hendrickson. 522.— Rivero. A. 95. 1957. 1961. p. p. Meta: MCZ 16269-71. Villavicencio. p. Hyla creolica Werner. p. 522. 1959. gularis Werner. — p.. 1944a. 1944a. 133.—Hunter and Murcia de V. p. labialis labialis The length of crus more than half the head-body length H. must await the receipt of fresh material before deciding on the status of the frogs Ruthven that forms (1922. Hyla servalina Werner.— Rivero. 72. 1944a. 1944. b 2. p. The three be distinguished as follows: Posterior face of the thigh uniformly pigmented or lightly speckled but not distinctly mottled. Tatarenda. p. 1957. 1899. Specimens Examined COLOMBIA Boyaca: Sutatenza. 10. 263— Dunn. 108. p. sabana near Bogota^. Boquer6n near Bogota^. 1959. believed neighborhood of Bogota). p. 1961. 302 (type locality. 1961. 1959. 1923. 3 (type locality. p. We platydactyla. Hyla 8. p. 1882a. p. p. mimetica). 463 (type locality. Chaco. 1962. 10. 469 (type of H.— Heimlich. 72. p. 1940a. p. De Luque and Hunter.133 (type of Hyla palpebrogranulata) Roque.— Boulenger. Nieden. p. 1957. 460. 72. Hyla wilsoniana. p. 522. 484 (type locality. p. a2 . NR BOLIVIA: PERU: labialis group 14. There are at present three vicarious forms in this group H. USNM 152089-97. and H.— Rivero. a K b may l . 1914. p. 1863b. labialis platydactyla . . Ex. p. 484 (type locality.— Stebbins and Henderickson. labialis krausi. Hyla vilsoniana Cope. p. 1957. labialis labialis Peters Plate 33d-p 1863. 1899. 522. p. 1923.— Dunn. 10. I. 1957.

nostrils as lateral as superior. roof of skull not exostosed. the upper jaw extending but little beyond lower. that of belly and coarsely granular. labialis hausi in having shorter and the length of the crus less than half the head-body length. no dermal appendage on heel. pal- membrane not reticulate. ColomVomerine teeth in two small. —A patagium which 2 88 frog of the labialis group with a well-developed is bright blue in life. Snout short.S. tongue one-half as wide as mouth Description. no projecting rudiment of a pollex. no ulnar ridge. the web on fourth toe reaching base of penultimate phalanx. interorbital distance slightly greater than width of upper eyelid and slightly greater than distance between nostrils. not prominent. glandular ridge encircling upper part of tympanum. Tympanum distinct. a rather narrow. disk of fourth toe covering about one-half the tympanic area. bia. platydactyla in having the groins and the posterior surfaces of the thighs uniform or lightly speckled but not distinctly This subspecies differs from Hyla legs mottled. its diameter equal to its distance from nostril. broadly rounded. Cundinamarca. just reaching to disk of third whioh covers about one-half the tympanic area. lying close together between the small. opening. Body pebral short and stout. no outer but a weak inner tarsal ridge. a distinct oval inner and a smaller but distinct rounded outer metatarsal tubercle. heel reaches tip of snout.252 NATIONAL MUSEUM BULLETIN U. Skin of upper parts smooth. fourth finger considerably longer than second. Diagnosis. thick. — USNM 95155. Skin of head not co-ossified with skull. a welldeveloped skinfold across chest. adult female. heels overlap conpatagium extends from the back of the upper distinct to the side of the body. its posterior border slightly free and very shallowly notched. I. the upper lip flaring out a little below it. Can thus rostralis moderately defined. rounded choanae. skin of throat and chest somewhat wrinkled and lower surface of thigh uniformly in appearance. no vocal sac. when hind siderably. . Fingers webbed at base. the web reaching base of antepenultimate phalanx of third finger. third and fifth toes subequal. from Bogota. about one-half the diameter of eye. Eye small. It differs from H. separated from each other by an interval equal to about their distance from eye. Toes slightly less than one-half webbed. knee and elbow overlap appreciably. rounded patches. not projecting. separated from eye by a distance equal to its own diameter. when hind leg is adpressed. arm legs are A bent at right angles to body. slightly rounded in profile. somewhat triangular when viewed from above. in postaxillary region about equal greatest width of head. when limbs are laid along the side. their distance from end of snout about one-half that from eye. loreal region nearly flat and slightly oblique.

not in pattern). where the dark gray of the dorsum merges gradually with the cadaverish gray of the venter. but it is impossible. Very frequently the green is not uniform. The black line may be lacking in some bright green individuals (types of gularis). except for a little darkening or lightening of the general tone. There is some ability for color change in individuals in life (in and most of the uniform specimens darken in preservation. 18. While the two individuals do seem in general characters to be they cannot. diffuse black pigment producing darker sides and a darker dorsum. Dr. After having collected and studied the form in life in Bogota and environs.. labialis or H. 1957. Two specimens in the British here referred to this form. femur. tibia. I. p. I have kept these different color forms alive for days and the individuals maintained their colors constantly.. width. in our opinion.. platydactyla. the vomerine teeth series seem to be shorter than in typical H. and hands.5 millimeters in head- perhaps related to the referred to either H. 55) report of the species from Santa Marta. I. the other is an adult female that measures 24. in the concealed surfaces and. Remarks. Likewise. labialis. the "servalina" spotting. — The most frequent paler below coloration and with blue is a rather uniform bright light green above. An occasional frog may not be any shade of green at all but a golden brown. body length. feet. to tell if it was pigmented in life. head length. black "servalina'" spotting. I. the two separated by a light dorsolateral band. inconspicuous small black specks may be seen. hand. 20. They do have a patagium.5. A combination of the striping.4 mm. with green and golden brown each forming irregular parts of the background produces a very gaudy frog indeed. 39. labialis or H.6 mm. in their faded condition. 11): Color in alcohol. 15 —COCHRAN mm. heel-to-toe. head mm. 30.8 mm. p. This was the color of the types of labialis and of vilsoniana. either uniform or with the lighter dorsolateral band.2 mm.. we can add nothing pertinent to the above discussion.. 15 mm. I.15. Dunn reported on the pattern variation as seen in labialis. of the two is . in large individuals. and the dorsum divided by a lighter and vaguer middorsal light line wider and more prominent posteriorly. but there are also very dark olive green individuals (almost black) and the black line does not show up in these either (types of creolica). be platydactyla. I.23-24) are now pale gray with black. These also may have the tone. appreciably smaller than adult females of H.4 Dimensions. Along the sides. A black line runs from the nostrils back over the tympanum to widen and fade out on the body. and we quote his discussion here (Dunn. Green individuals may be irregularly spotted with black on the whole dorsal surface (type of servalina). 48283-4) that formed We have seen the two specimens its (MZUM the basis of Ruthven's (1922. 12. One Museum (BM 1902. on the belly also.FROGS OF COLOMBIA AND GOIN 253 — Head and body. a few similar specks are present on the lateral margins of the shanks.. labialis group. — A rather dark gray frog without much pattern. One of the specimens seems to be immature.

it differs from H. 57487. 1900. Arracachal. arranged more or less in rows such as one finds in the "servalina" pattern. CNHM 81895.3. Cundinamarca. Tabio. ZSM 102/1937 (holotype). 152176-8. CJG 2290 (24). separated from each other by an interval about equal to their distance from eye.35-6. detailed report of oogenesis in this species. Tunja.23-4. Paramo de Sumapaz. Laguna de la Rivero. Cundinamarca. ANSP 25772.2.5 [1947.4. Villeta. USNM 152057-9. Colombia. 76071.S. nostrils as dorsal as lateral. MLS 260. 78310 (52). 92161 (14). CJM 1252 (4). (1962.3.6. Eye — . 573-583) recently published an excellent.5. Paramo 81893-4.2. Hyla vilsoniana krausi Hellmich. transverse series. Villavicencio. slightly projecting. Bogota. FM 287 (21). Hunter and Murcia de V. Aguadita. lying rather far apart between the small. de Verjon. Paramo. 95152-76. Guitarra. Vomerine teeth in two short. BM 1915. 68. CNHM Cundinamarca: BM USNM CNHM MZUM MCZ USNM 152083-7. Description. Specimens Examined COLOMBIA Laguna de Tota. 8 (type locality. rounded choanae.7. 152088. its posterior border free and very shallowly notched. Cruz Verde 152171-3. Hyla labialis krausi Hellmich Figure 24 1940. USNM 152034. Madrid. 95. Can thus rostralis well defined. from Laguna de la Guitarra.5-6. Hyla labialis krausi differs from H. p.6-14. Boy ac A: 20289. 1909. 152159-70. SUNHM Paramo Vijogual [20 km.4. Cota. north of Pajarito].23. Paramo de Sumapaz. 89423 (31).3. near Bogota.254 NATIONAL MUSEUM BULLETIN 288 U. 1919. 94958-66. Snout moderate. CNHM 1902. somewhat U-shaped when viewed from above. USNM USNM USNM BM Meta: 152174-5. labialis in having longer legs and the length of the crus more than half the head-body length. Paramo de Palacio. their distance from end of snout about equal that from eye. Colombia) — 1963a. loreal region slightly concave and somewhat oblique. broadly rounded.' A large frog of the labialis group with a well-developed patagium which is bright blue in life. pp. Choachf. 95872-921. JRT 34 unnumbered.14] (type of creolica). 57494.15. — Diagnosis.7. 81897. vermiculate spots on the back. 81892. 78302 (14). 1940a. ANSP 24335-9. broadly rounded in profile. the upper jaw extending but little beyond lower. CNHM 81896. Gutierrez. the upper lip flaring out a little below it. 9046-51.11. MZUM 89455. platydactyla in that the groins and posterior surfaces of the thighs range from uniform to lightly speckled but are never distinctly and brightly mottled. p. tongue three-fourths as wide as mouth opening.6.

Toes slightly the web on fourth and more than one-half webbed. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. just reaching to disk of third which covers about one-half the tympanic area. subequal. no tarsal ridge. its diameter greater than its distance from nostril. about three-fifths the diameter of eye. Hyla labialis krausi. separated from eye by a distance nearly equal to two-thirds its own diameter. holotype. Tympanum distinct. fourth finger considerably longer than second. heel reaches to nostril. knee and elbow overlap considerably. when hind leg is ad pressed. Fingers webbed at base. no ulnar ridge. Skin of upper parts smooth. a rather narrow glandular ridge passes above the tympanum. . palpebral membrane not reticulate. very prominent. no projecting rudiment of a pollex. A patagium extends from the back of the upper arm to the side of the body. when limbs are laid along the side. toe reaching the base of the penultimate phalanx. no dermal appendage on heel. when hind legs are bent at right angles to body. ZSM 102 (same size). in postaxillary region about equal greatest width of head.— —COCHRAN FROGS OF COLOMBIA AND GOIN 255 moderate. skin third fifth toes third the Figure 24. heels overlap greatly. which is slightly greater than distance between nostrils. disk of fourth toe covering about onetympanic area. interorbital distance about equal width of upper eyelid. Body not elongate.

302. roof of skull not exostosed. Meiida. p. Venezuela). 16276. Venezuela).5 mm. 26.6 mm. Laguna de Paramo de Sumapaz. p. 39. ZSM 102/1937 (type). 1.. Diagnosis.— Gorham. its pos- and unnotched. vocal sac apparent as wrinkles on the throat. The anterior and posterior surfaces of the thighs are lightly mottled but not nearly so heavily or conspicuously H. Remarks. the type. brownish gray below. pencil-like flecks and spots. p. 1923. 1961. p.—Lutz. All of the dorsal surfaces are marked with tiny. Hyla vilsoniana meridensis Rivero. Vomerine teeth in two short. Hyla platydactyla Boulenger.4 mm. Snout short. of throat and chest and lower surfaces 2 88 finely granular. 15. 131 (type locality. 50. as all the paratypes were destroyed during World War II. 262. 115. traces of a skinfold across chest. from Chita. Hellmich for the privilege of studying and illustrating the type specimen. tongue three- opening. p. krausi in that the posterior faces of the thighs are distinctly mottled. The ventral surfaces seem to be without pattern. nostrils nearly lateral.— 256 NATIONAL MUSEUM BULLETIN U. MCZ 16032-3. heel-to-toe 39. —A frog of the labialis group with a well-developed patag- ium which is bright blue in life. labialis platydactyla. 1961. Description— 92162. rounded when viewed from above. nearly transverse series lying fairly MZVM close together between the fourths as wide as mouth small. 1963a. is now represented by a single specimen. Head and body. 183 (type locality. 23 mm. — Dimensions. The type series of this form. 1963a. femur. labialis and H.S. 24.— Rivero. Dark — bluish gray above. p. rounded choanae. very broadly cordiform. I. Adult male. cadaverish. nearly truncate and very slightly sloping in profile. Nieden.. 1927. which is apparently valid. terior border free . We are greatly indebted to Dr. tibia. head width. the upper jaw extending but slightly beyond lower.2 mm. that of belly coarsely granular.. The nuptial pads on the thumbs are very dark gray.. Boyaca. Hyla labialis platydactyla differs from H. 17. Colombia. so as in — Specimens Examined COLOMBIA Cundinamaeca: Gutierrez. 1963. 1964a. of thighs uniformly and very finely granular. Skin of head not co-ossified with skull. 93. Merida. p. p.. head length. hand.5 mm. The three specimens from Gutierrez are the only ones we know of other than the type. 93. Hyla labialis platydactyla la Guitarra. p. 22. black. Boulenger Plate 34a-c 1905.5 mm. Color in alcohol..

a rather distinct skinfold across chest. just reaching to disk of third which covers about three-fourths the tympanic area. loreal region very slightly projecting. with a distinct pattern of vermicidations. Head and body.2 mm. roof of skull not exostosed. Fingers webbed at base. heels overlap appreciably. Tympanum very distinct. not prominent. Skin of head not co-ossified with skull. no projecting rudiment of a pollex. head length. — — — . The axillae. Toes slightly more than one-half webbed. in postaxillary region just a little narrower than greatest width of head. femur. heel-to-toe. their distance concave and nearly vertical. hand. no vocal sac. 39. belly. toe reaching nearly to tip of antepenultimate phalanx. a rather narrow glandular ridge passing above tympanum. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. nearly black. Skin of upper parts smooth. head width. palpebral membrane not reticulate. skin of throat finely granular.7 mm.. Dimensions.5 mm. The only question is whether the specimens we examined from Boyaca should be listed as typical laid along the side. Canthus rostralis very well defined. the no ulnar web on fourth ridge. heel reaches to nostril. tibia. disk of fourth toe covering about three-fourths the tympanic area. Remarks. no tarsal ridge. anterior and posterior faces of thighs. Color in alcohol. The tops of the arms and hands and the shanks and feet are bluishgray. its diameter equal to its distance from nostril. as the dorsum. ventral surfaces a dark cadaverish gray.. when hind legs are bent at right angles to body. no dermal appendage on heel. 19. Eye moderate.. and undersurfaces of shanks and feet very dark gray. about three-fifths the diameter of eye. A weak patagium extends from the back of the upper arm to the side of the body.. groins. that of chest..8 mm. fourth finger longer than second. Top of head and dorsum uniform bluish-gray.9 mm. There seems to be no doubt that the population of Hyla labialis in the northern portion of the Sierra Oriental of the Andes in Colombia and Venezuela is sharply different from the population of typical labialis around Bogota. 12. 29. interorbital distance one-third again the width of upper eyelid. separated from eye by a disslightly at all tance equal to two-thirds its own diameter.5 mm. which is slightly less than distance between nostrils. when hind leg is adpressed.. third and fifth toes subequal. the upper lip flaring out hardly below it. but on top of the thigh the blue-gray of the dorsum is restricted to a rather narrow stripe of ground color pigment that separates the pattern of vermiculations on the front of the thigh from that on the back. 13 mm. Adult female. and lower surface of thigh uniformly and more coarsely granular. Body not elongate. 21.FROGS OF COLOMBIA —COCHRAN AND GOIN 257 from end of snout about one-half that from eye. 10. when limbs are knee and elbow meet snugly.

2. 1923. b1 . Cali. —A small frog with well-developed patagium. however. near Merida.— Dunn. while the type of platydactyla. 120 (type locality. We have seen the type of platydactyla and that of meridensis. Specimens Examined COLOMBIA MZUM BM 92162 (19). 280.—Barbour and Loveridge. p.14] (type of platydactyla). . p. These Either belly or dorsum. Colombia).13. p. 2527 (type of meridensis). Merida. Chama. appears to have more of a "servalina" pattern (see under H.l.— Cochran. p.17 [1947. 60.— Marx. 1958. H. group of the Sierra Occidental likewise consists of three vicarious may be distinguished by the following key: this forms.S. 1952. rounded vomerine tooth patches and with well-developed patagia which are pink or red in life. VARIABILIS group Small to medium-sized tree frogs with small. 17. throat. Diagnosis. which occupies the Sierra Oriental. 434. 1904. 71114. Nieden. with a rather distinct pattern. belly imb2 maculate or lightly mottled H. in having the belly much more extensively mottled. and on the basis of an examination of these types we are calling the Boyaca specimens typical platydactyla. more distinct dorsal pattern extensive mottling on the undersurfaces. Hyla and bogerti. labialis). From variabilis is From columbiana and much more most it differs closely allied to columbiana in having a bogerti it differs in having a longer leg so that the extended heel reaches the anterior rather than the posterior margin of the eye. 1961. bogerti No distinct dorsal pattern and ventral surface nearly or completely ima2 a1 .6. a series of fresh material from the Venezuelan Andes is required before the limits of variation in platydactyla can be ascertained. p. Valle. 1944a. and chin.30. or both. As in the previous group. and belly generally rather heavily mottled in dark gray or black. columbiana maculate Hyla variabilis Boulenger Plate 34d-f 1896. 1929. 77. The type of meridensis is very similar to the Colombian specimens from Boyaca. Hyla variabilis Boulenger. and in having . although somewhat old and faded. VENEZUELA: USNM MCZ 15.— 258 U. ventral surfaces generally H. variabilis heavily mottled Dorsal pattern generally of rather distinct dorsolateral stripes. p. Boyaca: Chita. NATIONAL MUSEUM BULLETIN 288 platydactyla or whether they should be listed as intergrades. reduced webbing between the fingers. . p.— Peters. 262. Dorsal pattern of blotches or interrupted stripes.

. tongue three-fourths as wide as mouth opening. femur. Since much of the preserved material is old and somewhat faded. Canthus rostralis rounded. belly. loreal region slightly flattened. tibia. its diameter equal to its distance from nostril. third toe slightly longer than fifth. Color in alcohol. broadly cordiform. separated from each other by an interval equal to about two-thirds their distance from eye.6 mm. Toes about three-fourths webbed. Skin of upper parts smooth. 25. Body not elongate.2 mm. an oval inner and a smaller.. at 3. Dimensions. no rudiment of a pollex. knee and elbow overlap slightly. Valle. a male and two females. when hind leg is adpressed. rounded patches lying close together between the choanae. disk of fourth just covering the tympanic area. Tympanum distinct.200 feet. separated from eye by a distance slightly greater than its own diameter. nostrils more lateral than superior. 13. Colombia. no tarsal ridge. Wilmot A.. from Cali. not quite reaching to disk of third which covers the tympanic area. heel reaches almost to nostril. skin of chest. Head and body. the upper lip not flaring strongly. somewhat triangular in shape when viewed from above. when hind legs are bent at right angles to body. 12. the upper jaw not extending beyond lower. no dermal appendage on heel. slightly defined. a definite skinfold across the chest. head width. fourth finger considerably longer than second..3 mm. 7. in postaxillary region about equal to greatest width of head. heavy. not projecting.—COCHRAN FROGS OF COLOMBIA dorsal stripes (such as there rupted. about one-third the diameter of eye. Thornton. indistinct rounded outer metatarsal tubercle. 8. vocal sac apparently internal. hand. The adult females are rather dark brown above and heaviJy mottled below.8 mm.5 mm. which is relatively wide. A moderately developed fold of skin (patagium) extends from the back of the upper arm to the side of the body.. Valle. unfaded material based on three specimens. and lower femur uniformly and very finely granular. 8 mm. Eye moderate. On one of the females are quite distinct postocular dorsolateral Description. at Candeleria. near Cali.. heel-to-toe. above and below. 16. their distance from end of snout about one-third that from eye. — — . Vomerine teeth in two short. Fingers practically free of web. heels overlap moderately. slightly greater than distance between nostrils.8 mm. head length. somewhat truncate in profile. we present the following description of fresh. its posterior border slightly free and very faintly notched. in June 1962. Snout moderate. when limbs are laid along the side. collected by Dr. interorbital space about the width of upper eyelid. may be) on —USNM AND GOIN 259 the back that tend to be inter- 71115 (cotype). This specimen is now a faded uniform pale brown. Cauca Valley.

making shallow ponds. the dorsal surfaces of the limbs are without distinct pattern.7 10. on the anterior face of the thigh is a pale. and in the heavy vegetation in the drainage ditches along the variabilis Where water collects in .8 can be found breeding throughout the Cali savanna. but posterior to this the light of the dorsolateral stripe merges ventrally with the ventral ground color and thus the dorsobecomes obsolete. In this specimen the dorsal surfaces of both the front and hind limbs have scattered dark flecks. 151983 (?) 29. is The heavily and coarsely vermiculated with a black and white pattern. muddy brown. and. —In shown is a This is addition to sexual differences in pattern. although a hint of a paler postocular light stripe can be seen. including stripe A those of the thighs. —In life the portion of the ventral surface which is was bright orange and the dorsal surface was dark. is and belly entire undersurface of the chin. of the three fresh specimens described above. NATIONAL MUSEUM BULLETIN 288 light stripes that continue from the eye posteriorly to behind the shoulders. unpigmented band between the dorsal pigmented surface and the ventral pigmented surface. measurements. In both specimens the undersurfaces of the thighs are darkened but not particularly vermiculated. male collected with the above two females has the dorsal pattern very similar to theirs. there much noticeable difference in size.1 6 ( 29. There are faint little pencil-like dark flecks in the dark dorsal ground color. chest. in millimeters. low places in the savanna. whereas the male was light orange-brown dorsally and immaculate below with just a tinge of orange-yellow on the ventral surfaces.9 14. but there is no discernible pattern.) 260 U. Likewise.S. 8 14. In this male the entire undersurfaces. The other large female lacks the distinct dorsolateral light stripes. thus leaving a clear longitudinal on the anteroventral surface of the thigh. in both. Specimen USN M151984 UV 236 9 USNM Hyla Head-body (<?) 21. Color in life.7 15. the females being in the following larger. are immaculate. throat. 5 13. but with the postocular light stripes even paler than in either of them.8 Thigh Shank 9. pale in the preserved females Remarks. lateral stripe Ventrally the pattern of these two females quite striking. In the same specimen the top of the head from the nostrils to between the eyes is nearly as pale as the dorsolateral light stripes. although their ground color is lighter than the ground color of the dorsum. as if the individual had been sprinkled with black pepper.

Antioquia.65-81 [1947. 60251 MCZ (2). 118729-30. difficult to collect. — USNM 118731. 7665. 51269.15-31]. palpebral 337-262—70 18 . Canthus rostralis moderately well defined.2. not prominent. 152030-1 90601. Antioquia. 63897.13. Antioquia. near Cali. rounded patches. new species Figure 25 Holotype. MZUM CNHM USNM 75989. This new species is most closely allied to variabilis and columbiana. separated from each other by an interval nearly equal to their distance from eye. Vomerine teeth in two small. Specimens Examined COLOMBIA Value: CNHM BM MCZ 95. the upper jaw not extending appreciably beyond lower. ANSP 25773. ablis). from Pacora. rounded choanae. MLS 358.11. these frogs are they are —COCHRAN AND GOIN 261 although at times calling. 118243. UV 236. a pale patagium. loreal region slightly concave and slightly oblique. CNHM MZUM MCZ 24911-5. 71115 (all cotypes of Hyla vari46464. Antioquia. USNM CNHM MZUM MCZ USNM USNM from Yarumal.16. Nic6foro Maria. A small Hyla of the variabilis group with reduced webbing between the fingers. Paratypes. from Jeric6. their distance from end of snout about one-half that from eye. Antioquia. 24888-93 (+200 dupl. From columbiana it differs in its shorter leg and in having a distinct dorsal pattern. — Description of holotype. Candeleria. Antioquia. 15076-100. nearly truncate in profile. Snout short. CJG 2280 (8). 152021-9. from Medellin. broadly rounded. From the former it can be distinguished by having a shorter hind leg (heel reaching posterior margin rather than anterior margin of ey e )> by having the pattern generally of two fairly distinct dark stripes on the back instead of broken lines and blotches. and generally with — a pair of fairly well developed darker stripes down the back. near Cali. 7664. Cali. 71214. from near Medellin. Eye small. and by having less mottling below. Caldas. from La Ceja. nostrils more lateral than superior. its diameter equal to its distance from nostril. Cerro de La Herrara. ANSP 21029. 17098-101. hardly projecting. from Rio Porce. lying close together between the small. MZUM USNM USNM USNM Hyla bogerti. near Cali. the upper lip flaring out a little below it. 58908 (10). from San Pedro. Antioquia.). 56508 (38). 63884. its somewhat rounded when viewed from above. posterior border slightly free and unnotched. tongue two-thirds as wide as mouth opening. Colombia.—ANSP 25005. 151983-4. 2606. 137770-2. 92167. 78301 (17).FROGS OF COLOMBIA commonly heard roads. 3565. by Hno. Diagnosis. Antioquia. 71215 (2). 30567-9. collected at Medellin. 63883.

which is relatively are bent at right angles to body. when hind legs of upper reticulate. no ulnar ridge. reaching to disk of third which snugly covers the tympanic area. the web on fourth toe reaching base of penultimate phalanx. Toes more than one-half webbed. separated from eye by a distance "about equal to its own diameter. heels overlap slightly. Tympanum fairly distinct. when limbs are laid along the side. disk of fourth toe covering the tympanic area. A small . heel fails to reach posterior margin of eye. knee and elbow fail to meet. about one-half the diameter of eye. no projecting rudiment of a pollex. Figuke 25.262 U.S. holotype. no dermal appendage on heel. then continuing as a thin margin to base of disk. no tarsal ridge. web reaching base of antepenultimate phalanx of third finger. a distinct oval inner but no apparent outer metatarsal tubercle. interorbital distance greater eyelid. when hind leg is adpressed. membrane not NATIONAL MUSEUM BULLETIN 288 than width narrow and slightly less than distance between nostrils. third and fifth toes subequal.— Hyla bogerti. Body moderately elongate. Fingers with but a trace of web at base. in postaxillary region a little narrower than greatest width of head. USNM 118731 (X 2). fourth finger considerably longer than second.

throat. femur. There seems to be no discernible difference between the specimens from Medellin and those from San Pedro.. 8... In some specimens the pattern is quite indistinct. and 33. otherwise the ventral surfaces are immaculate. and there are no distinctive markings on the thighs and lower legs although they are pigmented.3 mm. in which they range from nearly immaculate to being quite heavily mottled on the chin. — Dimensions. and belly. Thus we find columbiana in Cauca.0.7 Color in alcohol.0. one of which originates back of eye on each side and continues posteriorly to just behind the sacral hump.8. This new species represents the northernmost form of a series of small frogs with well-developed patagia that range in the Occidental Andes of Colombia from Cauca to northern Antioquia. that of belly and lower surface of thigh uniformly and finely granular a rather distinct skinfold across chest.5. 31. 31. adult gravid female. In life. whether this is due entirely to natural variation or whether it might be due in part to preservation is not known.5 mm. and just posterior to the eye.6.5 millimeters. The most conspicuous element of the dorsal pattern is a pair of dark stripes.. tibia. 31. In only a few of the specimens where the pattern is discernible are the pair of longitudinal dark stripes on the back interrupted or incomplete. In most specimens. The can thai region on each side likewise is of this darker hue. 15.. 31. There are some fuscous flecks on chin and throat.. Head and body. mm.7 mm.5 mm.1.. no vocal sac present. 33. Skin of head not co-ossified with skull. A round dark spot of the same hue lies between the eyes.1 mm.9. In the preserved specimens the most striking variation Variation. hand. skin of throat and chest nearly smooth. —A pale brown frog with a dorsal pattern of darker brown. 31. heel-to-toe.5. on each side. 10.2 mm. 32. Skin upper parts smooth. Remarks. While in general the dorsal pattern is made up of two rather discrete dark stripes. columbiana has a patagium that ranges from flesh-colored to . Each of these stripes is about the width of the upper eyelid. is noted in the ventral markings. head width. Ten large adults have head-and-body lengths of 30. head length. 30. 10. a band of this darker brown continues through the tympanum to above the shoulder. when the limb is laid alongside the body the heel reaches only to the posterior margin of eye. a rather narrow glandular ridge encircling upper part of tympanum.3. FROGS OF COLOMBIA —COCHRAN AND GOIN 263 patagium extends from the back of upper arm to side of body. 31. There also is some variation in length of hind leg. but in a few it reaches to the anterior margin. roof of skull not of exostosed. and bogerti in — — Antioquia. 21. variabilis in Valle. 14. there is variation in this. The adults are rather uniform in size. reminiscent of the condition in variabilis.

While both groups do have well-developed patagia and seem to be ecological equivalents. 1922. the upper lip not flaring out below it. labialis alive. not projecting. transverse series. we do not imply that they are closely related. It from the former in lacking extensive mottling on the chin. We take pleasure in naming this species for Dr. Bogert. this chain of species seems to be the ecological equivalent of the three forms (H. and belly. lying between the small. p. rounded choanae. 164. Eye moddiffers — erate. and H. but in preservative it does not seem to be heavily pigmented. 77. —A small frog without a distinct pattern on the back. In a way. minuta but differ in lacking both the distinct pattern on the back and the transverse white line above the vent and across the base of the heels. for we are not at all sure this is the case. Hyla columbiana is most closely allied to variabilis and bogerti. with a well-developed patagium. Popayan. p. I. 41 (type locality. rounded. platydactyla) with brightly colored patagia that range from south to north in the Oriental Andes of Colombia. throat. labialis krausi.— Mertens. tongue one-half as wide as mouth opening. Preserved specimens sometimes closely resemble H.— Nieden. — Dunn. Charles M. Of these we have seen only H. Hyla columbiana Boettger Plate 35a-c 1892. their distance from end of snout about one-half that from eye. 1944a. Hyla columbiana Boettger. its posterior border slightly free and unnotched. loreal region nearly flat and nearly vertical.— Rivero. the upper jaw extending a little beyond lower. p. its patagium is bright blue in life rather than reddish or yellowish as seen in columbiana and variabilis. Colombia. Snout short. 1964a. I. teeth in two short.264 U. Popayan). 1923. its diameter greater than its dis- . rounded when viewed from above. separated from each other by an interval equal to about their distance from eye. H. NATIONAL MUSEUM BULLETIN 2 88 red and variabilis has one that is flesh-colored to yellowish. 265. not particularly prominent. whose kindness in lending material has contributed so much to the present work. Diagnosis. We have never seen a live specimen of bogerti. Colombia. and with reduced webbing between the fingers. Cauca. Canthus rostralis weakly defined. Cauca. labialis. MLS 192b. 1944. 303. and from the latter in having a longer leg (the extended heel reaching the anterior rather than posterior margin of eye) and in lacking a well-developed pattern on the back. I. rounded in profile. p. Vomerine Description. Hyla colombiana. nostrils as lateral as superior. p. so we do not know the color of its patagium. one of the other.S.

Toes slightly more than one-half webbed. —In Remarks.. heels overlap greatly. The ventral surfaces are immaculate and a pale dirty brown. the web on fourth toe extending as a margin to the base of the penultimate phalanx. head width.3 mm. 25. palpebral —COCHRAN membrane not AND GOLN 265 reticulate. heel reaches anterior margin of eye.. no tarsal ridge. skin of throat and chest smooth. When a pattern is present on the back of columbiana it is generally asymmetrical. just reaching to disk of third which just covers the tympanic area. vocal sac median. subgular. Head and body.3 mm. minuta but the absence of a superanal white line and distinct pattern on the back will distinguish it from that species on close inspection. Faint traces of crossbands are present on the shanks..2 mm. This blotch is rather oblong. no ulnar ridge. fourth considerably longer than second. that of belly and lower surface of thigh uniformly and very finely granular. 8. the proximal end of the antepenultimate phalanx of the penultimate of the second and fourth fingers.7 mm. when limbs are laid along the side. adult male. 8. 8. a rather narrow glandular ridge encircling upper part of tympanum. The thighs are lightly pigmented but without pattern. —A pale brown frog with a blotch of darker brown on the back. There is no evidence of the little horizontal white bar above the vent and across the heels that is so characteristic of H. hand. no projecting rudiment of a pollex. a distinct oval inner and a smaller somewhat indistinct rounded outer metatarsal tubercle.. heel-to-toe. Fingers webbed at base.5 mm. A deep notch on the posterior end gives the blotch a V-shaped appearance. about one-fourth the diameter of eye. Body not elongate. knee and elbow overlap considerably.- preservative this species looks superficially rather H. In some specimens no pattern is discernible on the back but in others dark chocolate brown blotches often can similar to . 11. disk of fourth toe covering about two-thirds the tympanic area. A small patagium extends from the back of the upper arm to the side of the body. which is about equal the distance between nostrils. in postaxillary region a little narrower than greatest width of head. 13.5 Color in alcohol.FROGS OF COLOMBIA tance from nostril. when hind legs are bent at right angles to body. Skin of upper parts smooth. separated from eye by a distance nearly equal to the web reaching its and the base of third finger own diameter. minuta. when hind leg is adpressed. with one end between the eyes and the other end anterior to the sacral hump. tibia. femur. Tympanum very indistinct. — 18 mm. Dimensions. interorbital distance half again as great as the width of upper eyelid.. no dermal appendage on heel. head length. Skin of head not co-ossified with skull.. roof of skull not exostosed. traces of a skinfold across chest. mm. third and fifth toes subequal.

ANSP 24402 is recorded as having been collected in the vicinity of Bogota. thick. El Tambo. this is broken up into individual blotches. 44109. columbiana is somewhat golden. Hyla bokermanni Goin Figure 26 1960. On July 6. 43933-80. In life. CNHM LOCALITY UNCERTAIN: ANSP 16. Quintana. 1961. however. the junior author had the pleasure of collecting topotypes of this species in company with Dr. CNHM 25677. . 192b. belly light H. Hyla bokermanni Goin. the ventral surface of this species is immaculate. scattered. 43982-91. 1960a.S. pepper-like flecks. 24402. in others the dorsal surfaces of the hind limbs are simply a moderate brown with faint. Brazil) p. Occasionally smaller discrete blotches occur on the posterior half of the back posterior to the sacral hump. the dorsal surface of H. such blotches are apt to be more prominent anteriorly and sometimes are fused to form a broad shield-like figure on the back. The patagium of the living individuals is red at night but fades to an orangish hue in daytime. 152146-51. Fred Medem and Dr. USNM MLS ANSP 25418. with the dorsal blotches a little darker. the data must be in error. 82005-7. parviceps that a?. 152081-2. but these are never so large. a grandson of the collector of the type series. however. Carlos Lehman. small a1 . CJG 2274-8. A small patagium and an oval white spot on the anterior face of the thigh is bordered in black. The following key will help identify the only two species of this group now known from Colombia: Small. but in very large females occasionally there are small but distinct vermiculations in dark brown scattered over the undersurface of the throat and belly. NATIONAL l^USEUM BULLETIN 288 be discerned on the light brown dorsal ground color. Most often. Generally. running from between the eyes to the region of the sacral hump. Tarauaoa. ANSP 44050. 266 U. 721 (type locality. and conspicuous markings on the thighs. When present. In some specimens the dorsal surfaces of the shanks and feet are crossed with about three fairly distinct dark crossbands. or prominent as they are in variabilis. Specimens Examined COLOMBIA Cauca: Popayan.. brightly fingers. parviceps group marked frogs with reduced webbing between the rounded vomerine teeth patches. 54719. belly dark H. Acre. since this specimen is certainly columbiana and the species is not known from anywhere in the eastern range of the Andes. bokermanni No patagium and without a distinct oval spot on the anterior face of each thigh.

not particularly prominent. Canthus rostralis moderately defined. Hyla bokermanni. and with a large. palpebral membrane not reticulate. rounded series. H. lying on a level with the posterior halves of the small. interorbital distance nearly twice the width of upper eyelid. Vomerine teeth in Figubb 26. considerably projecting. Eye moderate. broadly rounded. loreal region concave and nearly vertical. from winch it differs in being of smaller size and in having shorter legs and Diagnosis. a different pattern.— FROGS OF COLOMBIA — COCHRAN AND GOIN 267 — A small Hyla with reduced webbing between the fingers. truncate in profile. the upper jaw barely extending beyond lower. small. parviceps. oval light spot on the anterior face of the thigh that is surrounded by an ocelluslike dark band. their distance from end of snout about one-half that from eye. its posterior border slightly free and shallowly notched. CJG 2355 (X 2). the upper lip not flaring out strongly below it. from Rio Manso. rounded choanae. rounded patches of vomerine teeth. bokermanni is perhaps most closely allied to H. rounded when viewed from above. nostrils more lateral than superior. two small. Snout short. tongue three-fourths as wide as mouth opening. its diameter slightly greater than its distance from nostril. separated from each other by an interval equal to about three-fourths their distance from eye. Cordoba. . —CJG 2355. Colombia. which is about equal the distance between Description.

Skin of head not co-ossified with skull.5 mm. adult female. and continuing posteriorly as a conspicuous dorsolateral stripe to a point about halfway between the sacrum and groin. when hind legs are bent at right angles to body. femur. about one-fourth the diameter of eye. heels overlap greatly. Body not elongate. roof of skull not exostosed. 7 mm.2 mm. disk of fourth toe snugly covering the tympanic area. no projectnostrils. The anterior face of each thigh is conspicuously marked with a large oval white spot that is bordered in black. Dorsal ground color dark chocolate brown. tibia. 11. — Structurally and in pattern the series of fresh specimens from the Rio Manso varies somewhat from the type but whether this is due to the fact that the type died before preservation. On each side a narrow white mark extends downward and forward from eye to the edge of upper lip and another wider white band extends downward and backward from eye to the edge of upper lip.5 mm.. the web on fourth toe just reaching the base of the half penultimate phalanx. fourth finger longer than second.. when limbs are laid along the side. just reaching to disk of third which covers an area about one-third again as large as the tympanic area. NATIONAL MUSEUM BULLETIN 288 Tympanum small. hand. ridge. head length. 7.8 mm. The dorsal and posterior surfaces of thighs are dark but are marked by slightly paler. head width. 12. A narrow pale stripe passes along the canthus from naris to eye. 8. separated from eye by a distance nearly equal to its own diameter. Head and body. skin of throat and chest smooth. . or a combination of reasons cannot be determined. lighter blotches. — — The ventral surfaces are pale. the digital Variation. Toes slightly more than onewebbed. 12 mm. not markedly distinct. distinct traces of a skinfold across chest. ing rudiment of a poll ex. A patagium extends from the back of upper arm to the side of body. Color in alcohol. heel reaches almost to nostril. that of belly and lower surface of thigh uniformly granular. 24. Dimensions. no vocal sac. a rather narrow glandular ridge encircling upper part of tympanum. or to geographic variation. no ulnar ridge. third and fifth toes subequal... or that the type is an immature specimen. in postaxillary region a little narrower than greatest width of head.. rounded. Skin of upper parts smooth. continuing along the edge of the upper eyelid to behind the eye where it abruptly widens to about the greatest diameter of the tympanum. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. this series differs in having the snout more truncate in profile.8 mm. knee and elbow overlap considerably. At any rate.268 U.. no tarsal no dermal appendage on heel. Fingers slightly webbed at base. when hind leg is adpressed.S. heel-to-toe.

and in down having the tips of the heels pale. and the clear ventral surfaces. Remarks.FROGS OF COLOMBIA — COCHRAN AND GOIN 269 disks better developed. the oval white mark surrounded by dark on the anterior face of the thigh. minuta. the knee and elbow touching when the limbs are laid along the side. Dorsally the pattern ranges from dark chocolate brown with distinct dorsolateral stripes (like the type and the one described above) to a pale gray dorsum speckled with dusky in which the only semblance of a dorsolateral stripe is a narrow stripe that continues along the upper eyelid posteriorly to above the tympanum. The variation in pattern is striking. tells us that these frogs were breeding in a small pond in the woods after the first heavy rain in a week.) . bokermanni has the oval spot on the anterior face of the thigh and lacks the white line above the vent and on the heels which is so characteristic of H. CJG Acre. we feel it best to leave them as nominal species. 2355-67. luteoocellala Roux. This species is certainly similar to Hyla rondoniae. in lacking a white ventral to the naris. 2881 (type). larger size (the type had a head-and-body length mark on the of 17. which is described from El Mene.8 mm. Until such time as a comparison between specimens of these three nominal forms can be made. Venezuela. The largest specimen at hand has a head-and-body length of 25 millimeters. Tarauaca. who collected the Rio Manso series. a definite patagium. The variation in structure in the series from Rio Manso is not noteworthy. Specimens Examined COLOMBIA C6rdoba: BRAZIL: Rfo Manso. the white spots below the eye. the hind leg reaching to the nostril when adpressed instead of just to the eye. Medem. Falcon Province. In some specimens the dorsal pattern is a dark brown hour-glass shaped figure running from the eyes to the middle of the back. Dr. Both of these dark brown marks are on a background of light brown and make a pattern so reminiscent of Hyla minuta that the two could be easily confused except for the fact that H. followed by a broad arched blotch of dark brown. He said also that the species previously had been unknown to him. in having the dorsolateral light lip stripe stop abruptly anterior to the groin rather than turning and passing into the ventral light color of the venter. WCAB . — which Bokermann recently described from Rondonia (formerly Guapore) Territory in Brazil. and it may be very close to H. The only elements of pattern common to all the individuals are the light line on the can thus.

MLS 54. skin of throat and chest smooth. from end snout about one-half that from eye. and a distinct light patch on each side below the region of the sacral hump. web on third finger reaching distal end of antepenultimate phalanx on lateral side and proximal end on medial side. p. knee and elbow overlap considerably. broadly cordiform. a distinct oval inner and a smaller indistinct rounded outer metatarsal tubercle. brightly marked thighs. Colombia. considerably projecting. no patagium. p. a small tympanum. separated from eye by a distance equal to one-half its own diameter.. p. about one-third the diameter of eye. 273. Ecuador. no ulnar ridge. rounded series. Diagnosis. its diameter one-third again as great as its distance from nostril. Sarayacu) 1926. Eye large. no tarsal ridge. rounded vomerine tooth patches. No patagium.— Melin. Skin of upper parts smooth. 22.— Gorham. sepainterval equal to about their distance from eye. Tympanum small but distinct. heels overlap appreciably. 638. p. the upper jaw barely extending beyond lower. Body not elongate. which is about equal distance between nostrils. Snout short. 393 (type locality. — A small brown frog with reduced webbing on the fingers. — Description. 1884a. the web on fourth toe reaching the proximal end of the penultimate phalanx. 1963. the upper lip flaring out a little below it. 23. Fingers webbed at base. short. 270 U. fourth finger longer than second. interorbital distance about equal width of upper eyelid. just reaching to their distance rated from each other of by an disk of third which just covers the tympanic area. an indistinct narrow glandular ridge encircling upper part of tympanum. p. nostrils nearly entirely lateral. Hyla parviceps Boulenger. palpebral membrane not reticulate. no dermal appendage on heel. when limbs are laid along the side. very prominent. 1882a.— Nieden. from Ceilan. loreal region concave and a little oblique. NATIONAL MUSEUM BULLETIN 2 88 Hyla parviceps Boulenger Plate 35d-f 1882. lying close together between the small. third and fifth toes subequal. when hind leg is ad pressed. 1941. rounded choanae. its posterior border very slightly free and very shallowly notched. disk of fourth toe covering about two-thirds the tympanic area. west of Florencia. Toes slightly more than one-half webbed.S. tongue two-thirds as wide as mouth opening. heel reaches just beyond tip of snout. in postaxillary region a little narrower than greatest width of head. Caqueta. no projecting rudiment of a pollex. that of belly and lower surface of thigh uniformly and finely . truncate in profile. Can thus rostralis well defined. Vomerine teeth in two short. when hind legs are bent at right angles to body. rounded when viewed from above.

The specimen described above was directly compared with the type in the British Museum and structurally the two were found to be identical. 2511. BM 80.5. no vocal sac. roof of skull not exostosed.6 mm. head width. 15. and then terminates in a point just below the sacral hump. hand. Brazil (WCAB 2511) the Brazilian specimen was found to differ only in being a little more warty on top of the head and in having a small wart on the apex of each eyelid. The ventral surfaces. brightly marked patagia and with well-developed ulnar and tarsal ridges. Ceilan. BM 1889. Specimens Examined COLOMBIA Amazonas: Caqueta: BRAZIL: Rfo Apaporis. melanargyrea.. The few specimens we have seen of this distinctive little — — The main frog are rather uniform. 11. while the MLS specimen is uniformly dark underneath.. head length. west of Florencia. Remarks.1 mm.. 17. Only one form . MARMORATA group Moderate-sized tree frogs with well-developed.9.13.10. WCAB Sarayacu. including the undersides of hands and feet.6 mm. gravid adult female. Banos. giesleri) has yet to be determined.5 mm. like the type. MLS 54. tibia. is at present known from Colombia. has a white patch on the chest and anterior part of the belly.4 mm. variation is in the intensity of pigmentation rather than in pattern or structure.FROGS OF COLOMBIA — COCHRAN AND GOIN 271 granular. are rather heavily pigmented with dark gray..8 mm. When the described specimen was compared with a specimen from Acre. — Dimensions. A light brown frog with bright markings on the Color in alcohol. thighs. Skin of head not co-ossified with skull. traces of a skinfold across chest. becoming lighter as it rises. The true status of several nominal forms in this group (senicula. The top of head and dorsum are a nearly uniform light brownish except for a ramus of the ventral ground color on each side that starts upward just anterior to the groin. femur. 7. ECUADOR: MCZ 28058-9.12. 5.30. Head and body. 12. The type has a white blotch on the throat and chest.51] (type).3 mm.. 21.2. heel-to-toe. The thighs are dark in color with distinct yellowish blotches on the anterior and dorsal surfaces and a single yellow spot near the base of each thigh on the posterior surface. The Brazilian specimen.214 [1947. Acre. 7.

650. p. Can thus rostralis slightly defined loreal region flattened and nearly vertical. Body not elongate. more lateral than superior. disk of fourth toe not quite covering the tympanic area. 1802. 1820. 1961.— Peters. somewhat and with a trace cordiform. its diameter equal to its distance from nostril. 174. p. La Providencia. 1841. 1838. third and fifth toes subequal. Merrem. in postaxillary region a little narrower than greatest width of head. its posterior border very slightly short.— Liu. distinct ulnar and tarsal ridges. lying close together between the small. just reaching to disk of third which just snugly covers the tympanic area. the web on fourth toe reaching the distal third of the penultimate phalanx. a well-developed ulnar ridge. 35. 1931. from Rio Orteguaza. 29 (type locality. 71. tongue about one-half as wide as — mouth opening. web extending to the middle of the penultimate phalanx of third finger. Snout . 1955. p. 3. Tschudi. Lophopas marmoratus. brightly marked patagium and well-developed ulnar and tarsal ridges. when hind leg is adpressed. — Diagnosis. p. 571—Fitzinger. p. 1838. fourth finger but slightly longer than second. The combination of brightly marked patagium. p. a little projecting. p. — — — p. their distance from end of snout about one-half that from eye. rounded series. 1871. no dermal appendage on heel. separated from eye by a distance equal to one-half its own diameter. 104. 18S2a. "Habitat Suriname"). Vomerine teeth in two short.S. 1802. 1803. interorbital distance greater than width of upper eyelid.— Cochran. 22. 1952. palpebral membrane not reticulate. 391 Crawford. heel free of a notch. the upper lip barely flaring out below it. Tympanum very distinct. A brightly colored Hyla with a large. Colombia.— Gunther. Daudin. a distinct oval inner and a smaller but distinct rounded outer metatarsal tubercle. Toes nearly completely webbed. 1868. 174.272 U. Eye neither large nor particularly prominent. an exceptionally well-developed tarsal ridge. nostrils tiny. the tooth patches are slightly larger than the choanae. separated from each other by an interval nearly equal to their distance from eye. p.— Cope. the upper jaw extending but little beyond lower. 34. and conspicuously spotted belly should serve at a glance to distinguish Hyla marmorata from any other Colombian Hyla. rounded when viewed from above. p. NATIONAL MUSEUM BULLETIN 2 88 Hyla marmorata (Laurenti) Plate 36a-c 176S. Fingers webbed. 126.— Boulenger. 73. 403. MLS 55. which is about equal to distance between nostrils. .— Rivero. Bufo marmoratus Laurenti. p. 1858. no projecting rudiment of a pollex. p. — — Calamita marmorata.— Gorham. 1963. p. p. rounded choanae.— Lutz and Kloss. 1820. 1935. Dumeril and Bibron. Description. about three-fifths the diameter of eye. 34. Hyla marmorata. p. nearly truncate in profile. p. p. 1843. Caqueta. 111.

roof of skull not exostosed. throat. adult male. 17. Rio de Janeiro) as a subspecies of marmorata and made dasynota Giinther (type locality. A well-developed.FROGS OF COLOMBIA — COCHRAN AND GOLN 273 when limbs are laid along the side. the arms of which come together above the shoulders. Skin of upper parts glandular. the patagium. rounded.3]) but we have not seen enough material . 21. Head and body. 127) considered reaches to tip of snout.5 mm. In the middle of the back the base of the Y-shaped mark is very broad and contains a light spot of the paler ground color. Rivero (1926. 26. and belly. At least three other nominal forms have been assigned to close relationship with Hyla marmorata. head width. The specimens from just east of the Andes in Colombia and Ecuador seem remarkably uniform. and the webbing on hands and feet are bright yellow. femur. hands (except the webbing). the undersides of forearms.9 mm..3 [1947. This inconspicuous dorsal figure starts at the posterior margin of each eye as a sort of Y-shaped mark. 35. A strikingly marked frog. The yellow of the chin. p. Remarks. 11. narrow. — — — melanargyrea Cope (type locality.11. A large.5 mm. a well-developed skinfold across chest.31. Brazil) a synonym of senicula. knee and elbow overlap considerably.7 mm. Corcovada. 11.. when hind legs are bent at right angles to body.. posterior. shanks. the base of the "Y" continues posteriorly to terminate between the sacral hump and the groin. Dimensions. that of belly and lower surface of thigh uniformly and more coarsely granular.5 mm. the anterior. the underside of ulnar and tarsal ridges. Color in alcohol. throat. tibia. The dorsal ground color is very dark brown with a nearly imperceptible dorsal figure of darker brown. belly. hand. and thighs is thickly covered with very distinct. dark above and patterned below. discrete. p. vocal sac hardly apparent. heels overlap considerably. The entire surface of the chin. Cochran (1955. and somewhat roughened with several well-developed warty protuberances on each side in the region above the posterior termination of the flare of the patagium. head length. and feet (except the webbing) are black. and ventral surfaces of thighs. On each side the patagium continues to the groin as a thin fold of skin. 174) placed senicula Cope (type locality. glandular ridge encircling upper part of tympanum. Skin of head not coossified with skull.5 mm. skin of throat and chest very finely granular. heel-to-toe.16. The undersides of forearms.. We recently studied a co type of melanargyrea (ANSP 11216) and the type of dasynota (BM 68.. Mato Grosso.8 mm. coal-black spots that range from one-half millimeter to two millimeters in diameter. well-developed patagium extends from back of upper arm to side of body.2. Brazil) as a subspecies of marmorata separate from senicula.. 10.

Rio Llushin. ECUADOR: 1 18. senicula. EPN 2601. EPN 1485. Among the Colombian frogs. Specimens Examined COLOMBIA CaquetX: La Providencia on Rio Orteguaza.—Stuart. The so status PHLEBODES group and relationships common in South America is of the numerous "small yellow frogs" impossible to determine at the present time. From third the diameter of eye). "Ecuador. 274 U. and we feel certain that intensive and extensive studies of these frogs throughout northern South America will bring about a rearrangement of these groupings. cabeceras del Rio Bobonaza. Hyla phlebodes Stejneger Figure 27 1906. 888. Hyla phlebodes is probably more closely allied to microcephala. p. 1923. p. north of Arapicos. Diagnosis. We set off the phlebodes group because it seems to us to be more different from the species in the microcephala group than the species in that group are from one another. 817 (type locality. although we do agree that marmorata. EPN 1812-4.S. and with the dorsal pattern consisting of a narrow bar between the eyes and an hour-glass shaped mark on the back that is made up of narrow dark lines. ran.— Taylor. little or no pigmentation . 1952. EPN 1431. MCZ 4776. San Jose Viejo de Sumaco. mathiassoni.— Coch- Nieden. Rio Napo. NATIONAL MUSEUM BULLETIN 288 from intervening localities to be dogmatic about the status of the several nominal forms. and riveroi. 1935. and melanargyrea seem to be different and that dasynota is probably a direct synonym of senicula. Alto Napo. H. The species in both of these groups seem to differ trenchantly from Hyla minuta. 57. MLS 55. cabeceras del Rio Capanuari. —A —Stuart. Costa Rica).— . Hyla underwoodi. San Carlos. JAP 3729. Only for the sake of convenience do we now recognize three groups of these frogs in Colombia. Napo-Pastaza Province. 39 (part) small. and it differs from frogs in the microcephala group in having a dorsal pattern consisting of a bar between the eyes and an hour-glass figure on the back. 1961. 1935. Hyla phlebodes Stejneger. EPN 2510-2. 253. yellow frog with a small tympanum (one- on thighs. p. p. p. It differs from microcephala in having an hour-glass-shaped pattern rather than longitudinal stripes and in having a rather distinct interocular bar." OV 5933. phlebodes differs from minuta in lacking the white bar above the vent and on the heels. 1935. Rio Villano. 39. p.

rather rounded in profile. its posterior border slightly free and very shallowly notched. nostrils more lateral than superior. very slightly projecting. the upper jaw extending slightly beyond lower. separated from each other by an interval equal to about their distance from eye. 81856. somewhat triangular when viewed from above. their distance from end of snout about one-half that from eye. CNHM 81856 (X 3). the upper lip not flaring out strongly Figure 27 — Hyla phlebodes. Snout short. Choc6. Finally. Vomerine teeth in two small. tongue two-thirds as wide as mouth opening. loreal region nearly flat and nearly vertical. rounded patches. from Andagoya. Can thus rostralis moderately well defined. Colombia. rounded choanae.FROGS OF COLOMBIA —COCHRAN AND GOIN 275 it can be distinguished by the presence of a dorsal pattern and the absence of conspicuous longitudinal dark lymph glands which run from the region of the shoulder to above the groin on each side. lying between the small. broadly rounded. —CNHM Description. . it differs from riveroi in lacking a pattern of dark bars that tend either to be arranged transversely across the back or in mathiassoni scattered blotches.

7 mm.. which is in turn about equal the distance between nostrils. no pattern is discernible. One such line forms an interocular bar. fourth finger slightly longer than second. Tympanum small but distinct. femur. adult male.2 mm. knee and elbow overlap considerably. palpebral membrane not reticulate. no tarsal ridge.2 mm. hand. The pattern consists of narrow.. A very weakly developed patagium extends from the back of upper arm to side of body. Skin of upper parts smooth.3 mm. 7. 11. — . no dermal appendage on heel. head width. like the dorsum. 6. separated from eye by a distance equal to about one-half its own diameter.. A pale. disk of fourth toe just covering the tympanic area. 21 mm. skin of throat and chest smooth. A rather distinct cruciform mark on the — back extends from just behind each eye to a point just anterior to the sacral hump on each side. Color in alcohol. heels overlap greatly. 7. Body not elongate. the five specimens from Colombia before us are rather uniform. no ulnar ridge. Fingers webbed at base. Variation. third and fifth toes subequal. just reaching to disk of third which is slightly greater than the tympanic area. about one-fourth the diameter of eye. heel-totoe. when hind legs are bent at right angles to body.S. in postaxillary region a little narrower than greatest width of head. 9 mm. There are very faint traces of darker crossbars on the shanks. Eye moderate but rather prominent. is a very light tannish brown. dark brown pencil-like lines on the dorsum. when limbs are laid along the side. Other than these marks. When the male from the Maracaibo Basin is compared with the one from the Pacific drainage. no projecting rudiment of a pollex. fourth toe about twice as wide at base as other toes. head length. 15. Head and body length. tibia. vocal sac large. the web reaching to distal end of antepenultimate phalanx of third finger. The venter. interorbital distance about equal to width of upper eyelid. median. NATIONAL MUSEUM BULLETIN 288 below it. very light brown frog. Skin of head not co-ossified with skull. no apparent traces of a skinfold across chest. its diameter greater than its distance from nostril.276 U.7 mm. a distinct oval inner and a smaller indistinct rounded outer metatarsal tubercle. heel reaches to between eye and nostril. that of belly and lower surface of thigh uniformly and very finely granular. roof of skull not exostosed.. but this may be due to differences in preservation. subgular. when hind leg is adpressed. no discernible significant variation can be noted. Toes slightly more than one-half webbed. Except for the degree of fading.. a rather narrow glandular ridge encircling upper part of tympanum. — Dimensions. MLS 119) seem to have brighter markings than the females. Two males (CNHM 81856.. the web on fourth toe reaching the distal end of antepenultimate phalanx.

89. underwoodi in London and agree with Taylor that Stuart (1935. 298. USNM 152116-7. Mato Grosso). 289. p. 1912. 1962. p.— Mertens. pp. Lutz.— Nieden. 1927. 29310-2. MCZ 2463 (2). 30. pp.— Miranda-Ribeiro. p. p. 1930. 34. 1959.— Miranda-Ribeiro. p. p. 1887. Janeiro). p. 1887a. 1872b. MEXICO: Yucatan. 1954. 1961. 1926. 1961b. 680 (type locality. 212 (type locality. bivittata — 1946. marshes in Rio and neighboring states). 1963.— B. 7. 122. Baumann. 1933. locality. 1926. 1960. near Porto Alegre. 417. 161. There is only one species known from Colombia. 1961. 496. 29224-5. p. near Rio de Janeiro). 6. p. Hyla yallens A. —B. 1957. Hyla emrichi Mertens. 1954. USNM 29970 (type). p. 471. 1959. 163.— De Witte.— Gorham. 123. 1888b. 1952. 22. Selva. p. 456.— Nieden. 46 (type locality. 1954. p. 1923. 227.— Myers. Astillero. 301. p. 120 Gans. Rivero. 1929. 1888. 1 (type locality.— B. 161. 287. 1925. p. Hyla suturata Miranda-Ribeiro. pp.— Miranda-Ribeiro. 1882a. 465. Turrialba.— B. p. 1912. Chapada. 1926. 1934b. Santa Catarina). Boulenger. 389. Hyla minuta Peters.— Cochran. 1955. MLS Andagoya.—B. Montserrat. p. Lutz. 163. 81. p. Lutz. 174. p. MCZ 28933-7. Boulenger. p. p. 161. 119. 162.— Cochran.— Baumann. Hyla 1912. Teres6polis.— Nieden. 13. Stebbins and Hendrickson. p. 1928. p. 1954. p.— A. p. 133. p. p. 287. with a modified hour-glass shaped figure on the back and with a transverse white line above the vent 337-262—70 19 . Lutz. Hyla bisittata. 57. p. 93 (type Lutz. NICARAGUA: 1.5 m.— Myers.—Bokermann. p. 294. p. — p. p. Lages.—Beebe. 1954. 1945. 39) erred in synonymizing phlebodes with underwoodi. Baumann.— Gallardo. p. 14. Lutz. p. 1912.— Cei and Reig. A small Hyla with reduced webs between the fingers and reduced pattern on the thighs. 12. Norte db Santander: La COSTA RICA: San Carlos.— Mertens. 31. Small. Hyla velata Cope. 57. 1963. 1926. pale to yellowish frogs with a dark saddle on the back and a transverse white line above the vent and on each heel. 11.— B. Rio Grande do Sul). minuta group 19. 158. 1961. 1958. MCZ 119. p. — — 91. 162. 1891a. 1923. 17-21— Gin6s. pp. 526. p. Rio de p. 1954. Lutz. Hyla minuta Peters Plate 36d-p 1872. p. figs. Chichen Itza. — Diagnosis. p.— FROGS OF COLOMBIA —COCHRAN AND GOIN 277 — Remarks. p. p. 1890. 1961. 163. p. Specimens Examined COLOMBIA Choc6: CNHM 81856. p. p. p. 1923.— Parker. p. 326. 1926. 1888c. We have studied the type of Hyla phlebodes in Washington and the types of H. north of Esteli. USNM 152152. 82.— Goin. 188 (type locality. Lutz.

hand. heel reaches tip of snout. third and fifth toes subequal. tibia.6 mm. its posterior border free and very shallowly notched. 8 mm.S. Description. fourth finger longer than second. no dermal appendage on heel.2 mm. head length. Toes slightly more than one-half webbed. Canthus rostralis slightly denned. . when hind legs are bent at right angles to body. which is about equal the distance between nostrils. Skin of upper parts smooth. Eye moderate. head width. that of belly and lower surface of thigh uniformly and very finely granular. Tympanum inconspicuous but distinct. slightly projecting. separated from each other by an interval equal to their distance from eye. when CNHM hind leg is adpressed. nostrils more lateral than superior. no tarsal ridge. Body not elongate.. Fingers webbed at base. adult female. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Colombia. Dimensions. when limbs are laid along the side. 12 mm. roof of skull not exostosed. a rather narrow indistinct glandular ridge encircling upper part of tympanum. about one-fourth the diameter of eye. no vocal sac. 8. 14 mm. heel-to-toe. 8. the web on fourth toe reaching base of penultimate phalanx. NATIONAL MUSEUM BULLETIN that coincides with white lines on the heels when 2 88 the legs are flexed. rounded choanae. 20 mm. the upper jaw extending but slightly beyond lower. heels overlap considerably.— 81764.. no ulnar ridge. minuta may be distinguished from members of both the leucophyllata and microcephala groups (the other small Colombian hylas with reduced pattern on the thighs) by the distinct transverse white line above the vent and on the heels. the upper lip flaring out but slightly below it. interorbital distance slightly greater than width of upper eyelid. 24.Vomerine teeth in two small rounded patches lying between the small.3 mm. Meta. its diameter slightly greater than its distance from nostril. loreal region nearly flat and nearly vertical.. knee and elbow overlap slightly. slightly rounded in profile. skin of throat and chest smooth. separated from eye by a distance nearly equal to one-half its own diameter. in postaxillary region about equal the greatest width of head. femur. Snout short. the web reaching the distal end of antepenultimate phalanx of third finger and to middle of penultimate phalanx of second and fourth fingers.. broadly cordiform.278 U.— Head and body. their distance from end of snout about one-half that from eye. Villavicencio. not quite reaching to disk of third which just covers the tympanic area... traces of a skinfold across chest. No apparent patagium. H. no projecting rudiment of a pollex. Skin of head not co-ossified with skull. palpebral membrane not reticulate. disk of fourth toe covering about three-fourths the tympanic area. tongue three-fourths as wide as mouth opening. somewhat rounded when viewed from above.

and part of may be correlated with geography. lack the white border to the dorsal pattern often seen in southern specimens. Three fairly distinct crossbars occur on each shank. pi. 2. and they (BM 6.Ribeiro. legs are flexed Variation. the sacral contiguous with similar lines across the bases of the heels when the and the heels held adjacent to the vent. its is range: The type locality of minuta is Rio de Janeiro. The other four specimens differ in that the main portions of the hour-glass figure between the eyes and across the back are not connected by the narrow longitudinal band of dark pigment. minuta in its several forms. The thighs have scattered. e-h). the only person who seems to have studied specimens in life from both the north (Belem) and the south (Sao Paulo). less conspicuous blotch occurs on the dorsal surface of the tip of snout. Padre Francisco Silverio Pereira. and suturata Miranda. As is characteristic of this species. .8. There is a single rather broad band across the back about halfway between the sacral humps and the tip of the urostyle. H. In three same as that in by Cochran (1955. In this specimen the consists an interocular of directed process that broadens out above the shoulders and divides arms of which extend nearly to the humps. —A AND GOIN 279 brown frog with dark brown main blotch on the back bar which bears a median posteriorly rather light blotches on the back. The ventral surfaces are immaculate. A smaller.9. Santa Catarina. The following additional names have been proposed: velata Cope.13. Mato Gross o. thus leaving a dark band between the eyes and another of seven specimens the dorsal figure is essentially the the specimen figured here and also in the specimen figured inverted U-shaped figure across the middle of the back. bivittata Boulenger. 1911. usually complete but sometimes with the anterior and posterior parts separate.83]). we here take the oppor- tunity of bringing the record up to date without trying to make decisions that should be withheld until the species as a whole studied throughout 1. emrichi Mertens. 1934. Rio de Janeiro. The dorsal pattern in the Rio population seems to be the hour-glass pattern. Rio Grande do Sul.6 [1947.—COCHRAN FROGS OF COLOMBIA Color in alcohol. The specimens from Colombia and Trinidad seem to be identical. Rio de Janeiro. pattern Lutz. and usually but not invariably with the dorsal pattern outlined in white. The pattern is quite variable in specimens in southern Brazil. informs us that the voice is distinctly different in the two populations. — Since there has been considerable confusion regarding the status of H. goughi Boulenger described from Trinidad is actually minuta the variation 4. minuta occurs naturally on Trinidad (Parker. cotype of H. 123). 3. now faded to a dirty brownish gray. 7. minute pigment spots but have no pattern. A 5.2. Remarks. while two less-distinct ones occur on each forearm. 9. a narrow white horizontal line above the vent is into a broad ypsiloid blotch. at —Most hand involve of the variations in the few Colombian specimens differences in the shape of the dorsal figure. p.

there seems to be no doubt that Hyla minuta is a species. Dorsum mottled with dark blotches. CJG 2383. 1959.83] 20.26 [1947.6 [1947. p. longitudinal dorsal stripes. p. and Mato 8. 1923. sacs b2. no conspicuous subdermal lymph b1 a 1 . 2336 (10). small. 1964a.7.2. 1960a. SURINAME: Onverwacht. Hyla under woodi. Thus. . and while the dorsal pigmentation in riveroi may occasionally be arranged in longitudinal rows.13. 1922.2. Until that day.— Nieden. Rio de Janeiro. bivittata) BM USNM 152108-11 96197-9 (cotypes of H. narrow. A.12. NATIONAL MUSEUM BULLETIN 288 Mr. Lages. 72. — Ruthven.2.— Rivero. that is in need of detailed study. goughi). 413. 484. 1960. reduced webbing rounded vomerine tooth patches. p. —A small yellow frog with a pattern of four dark.8. rather conspicuous subdermal lymph riveroi sacs. there are never more than two such rows and they are curved inward like a pair of reversed parentheses. MICROCEPHALA gTOUp with small tympani. riveroi of . 1931. H. 1901. p. 58. Diagnosis. p. but its dorsal pattern will distinguish it at once from these two species. Dorsal pattern of four narrow. USNM 88. Specimens Examined COLOMBIA Meta: Villavicencio. Dunn. p. The absence of an hour-glass pattern on the back will distinguish microcephala from phlebodes. 524. 257. 55. pallens). Bom BRAZIL: H. and without conspicuous markings on the thighs. Werner C. we must once again say maneat. p. H. Santa Catarina.— Stebbins and Hendrickson. H. mathiassoni Hyla microcephala Cope Figure 28 1886. yellowish between the frogs fingers. Hyla microcephala Cope. like most others in South America. microcephala apparently is closely related to mathiassoni and Colombia. 81764-6. mountain streams of ChiriquI. more-or-less distinct longitudinal dorsal stripes H. mathiassoni has no dorsal pattern. TRINIDAD: BM 1911.9. a*.S. The three species now known from Colombia may be differentiated as follows: Small.— 280 U. longitudinal stripes. 1922.— Gunther. 302. 265. p. p. Dorsum unmarked. Bokermann informs us that he cannot distinguish between the voices of the populations in Sao Paulo. ) (. 1933. (cotype of H. Grosso. 281 (type locality.— Fouquette. microcephala Dorsal pattern not of dorsal. Districto Federal. of CJG CNHM Successo. H. Panama). p.86-91](cotypes .

somewhat rounded when viewed from above. separated from each other by an interval equal to about their distance lower. broadly rounded. the upper lip barely flaring out below it. Eye relatively large and prominent. rounded choanae. lid. USNM 118248 (X 3). Description. between Tympanum its distance from interorbital distance equal nostrils and both greater than width of upper very distinct. the upper jaw extending appreciably beyond more than superior. about one-half that from eye. very slightly projecting. Magdalena. loreal region flat and nearly vertical.FROGS OF COLOMBIA — COCHRAN AND GOIN 281 — USNM 118248. slightly pointed in profile. the black choroid coat showing through from the translucent upper nostril.— Hyla microcephala. from eight miles northeast of Saiaza. Curumani. Can thus rostralis rather well-defined. its posterior border slightly free and very shallowly notched. lying close together between the small. palpebral to distance eyelid. its diameter greater than membrane not reticulate. about one-fourth the diameter of Figure 28. nostrils their distance lateral from end of snout eye. Vomerine teeth in two small. rounded patches. . Snout short. Colombia. tongue three-fourths as wide as mouth opening.

that of belly and lower surface of thigh uniformly and finely granular. Fingers when hind leg is adpressed. 6. 7. no ulnar ridge. third and fifth toes subequal.5 mm. The thighs are immaculate but there are scattered pigment cells on top of arms. heel-to-toe. thin. 11. hand. no dermal appendage on heel.. For from around the Fundaci6n-Aracataca region in Magdalena diverge Variation. Skin of head not co-ossified with skull. and each stripe continues back to above the tip of urostyle. The ventral — surfaces are immaculate. Skin of upper parts smooth. no outer but a low inner tarsal ridge. —We can discern geographic variation worthy of note in specimens example. separated diameter. knee and elbow overlap considerably. and feet. 17 mm. no projecting rudiment of a poll ex.6 mm. adult male. Toes more than one-half webbed.4 mm. A pale creamy yellow with longitudinal stripes on Color in alcohol. head width. skin of throat and chest smooth. head length. to tip of snout. pass above the tympani. the the Colombian specimens of this species. fourth finger considerably longer than second. A median pair of pale brown stripes originate on inner margin of upper eyelids. tibia.8 mm. NATIONAL MUSEUM BULLETIN 2 88 from eye by a distance equal to one-half its own webbed at base. traces of a skinfold across chest. 12. heels overlap greatly. shanks. in postaxillary region a little narrower than greateye. Head and body. and heavily folded.3 mm.282 U.5 mm. The sides of the body are slightly darker than the dorsum. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle.S. body. — Dimensions.. Another pair of stripes originate between the eyes and pass directly posteriorly. and extend along the sides nearly to the groins.. the dorsum. from the typical pattern of Hyla microcephala (which consists of four longitudinal dark stripes) as follows: A narrow dark stripe originates on the tip of the snout and passes posteriorly through the eye on each side and then passes straight back above the tympanum and shoulder to terminate in the region of the groin. rather narrow glandular ridge passes above the tympanum. a low. when hind legs are bent at right angles to est width of head. 23. the web on fourth toe reaching base of penultimate phalanx.. disk of fourth toe practically covering the tympanic area. reaching to disk of third which just covers the tympanic area. femur.. heel reaches almost when limbs are laid along the side. A patagium extends from the back of the upper arm to the side of the body. being separated by a .. Body not elongate. 7. web on third finger reaching middle of antepenultimate phalanx on lateral side and the base of antepenultimate phalanx on medial side. and the pigmentation where the dorsal and lateral side colors meet forms brownish stripes that originate at the posterior corners of the eyes. vocal sac subgular. roof of skull not exostosed.

are smaller in size (19 to 21. on the east bank of the Rio Magdalena in Cundinamarca opposite the state of Tolima. CJG 2444-5. something never seen in Colombian specimens from other localities.5 and 28. ANSP 19755-7. 136) points out that true misera ranges westward to the vicinity of Sierra de Santa Marta where it seems to blend with microcephala. on Rio Magdalena. Remarks.). and in two specimens the dorsal pattern between the lateral stripes is merely a series of blotches. Another series of specimens of Colombian microcephala showing divergence from the typical individual is from Girardot. MZUM 90168 (2). In all of these there is a marked tendency for the middorsal pair of stripes to break up into blotches.3 mm.7 km. USNM 152143. CJG 2457 (20). far and — Specimens Examined COLOMBIA Atlantico: Palmar de Varela. Told Viejo. Sincelejo-Tolu. 74919. 74918 (2). in head-body length) than usual. Curumani. Fundaci6n. Two specimens (MZUM 90168) from Curumani are likewise larger (24. ANSP 25061. 2459.23. Rivero (1961.) also show a tendency for the dorsal pattern to become modified. In one of the largest specimens from Aracataca (ANSP 19755. 26. and one is not so definitely striped as is usual for Colombian individuals.28]) and while we believe that it is certainly close to if not identical with H. microcephala we do not believe we can make a definite synonymic assignment without seeing ample fresh material from the type localities of both forms. JRT 390-3. Since we have not seen sufficient material from critical areas to of longitudinal demonstrate intergradation. BolIvar: Near Gambote. up the Magdalena Valley. CJG CNHM Cundinamarca: Girardot. MZUM 48281-2. Specimens in a series (JRT 604-12) from this locality.11. pp. 135. southeast of Bucaramanga. JRT 604-12.9 mm. and terminate above the tip of urostyle.2. the dark stripe on each side from tip of snout to groin is present but the rest of the dorsum and the top of the head and snout are marked with vermiculations which show hardly a sign arrangement but which do form an indistinct interocular bar.32-3 [1947. USNM 118247-8. It seems probable that the above-mentioned specimens simply show the influence of Hyla misera from the east. Santander: 4.2 mm. Choc6: Atrato.FROGS OF COLOMBIA —COCHRAN AND GOJN 283 distance equal to the interocular space. El Banco. with no middorsal stripes discernible. Magdalena: Aracataca. We have examined the type of Hyla underwoodi (BM 94. Rio Pechelin. .15. we presently consider all the material from northeastern Colombia that we have seen as microcephala and forego the use of the trinomial.

Can thus rostralis well defined. —CM — Goin and Layne. ANSP precise locality: USNM PANAMA: Hyla new riveroi. The —A tiny yellow frog allied to Hyla microcephala. rounded choanae. and the dorsal pattern of crossbars or blotches rather than an hour glass-shaped figure distinguishes it from H. fourth finger considerably longer than second. 120. disk of fourth toe covering about one-third the tympanic area a distinct oval inner and beyond lower. Colombia. Ama154 (2). Espinal. Meta. slightly rounded in profile.S. Toes slightly more than one-half webbed. Ross Allen in 1956. loreal region concave and nearly vertical. Fingers webbed at base. collected at Leticia. palpebral membrane not reticulate. third and fifth toes subequal.—CJG 2331-2. about one-fourth the diameter of eye. no ulnar ridge. Amazonas. its posterior border Snout short. 284 U. Sabana de San Juan der Arama. their distance from end of snout about one-half that from eye. El Real. (12). above. mathiassoni. interorbital distance slightly greater than width of upper eyelid. web on fourth toe reaching distal end of penultimate phalanx. Tympanum very distinct. Mariquita. broadly rounded. 146256-7. rounded series. which is relatively wide and about equal the distance between nostrils. its diameter half again as great as its distance from nostril. 8557 (4). Colombia. Eye large. Colombia. Tolima: No NATIONAL MUSEUM BULLETIN 2 88 CNHM 81822-3. from Acre. microcephala. CM 1958. rounded when viewed from slightly notched. misera Goin. 104. MCZ 15608. the mottled dorsal pattern and the lack of conspicuous subdermal lymph sacs in the dorsolateral region distinguish it from H. p. . separated from each other by an interval equal to about their distance from eye. by E. nostrils lateral . lack of parallel stripes on the back distinguishes Hyla riveroi from H. 37433. the upper lip flaring out below it only slightly. separated from eye by a distance about equal to one-third its own diameter. tongue threefourths as wide as free and mouth opening. Paratypes. very prominent. species Figure 29 Hyla 1958. p. Holotype. — Description of holotype. Darien. just reaching to disk of third which covers about three-fourths the tympanic area no projecting rudiment of a pollex. 19749. 2335. UF 8556 (6). 36085 zonas. phlebodes. 140567-8. MZUM USNM 125428-9. from Leticia. Vomerine teeth in two short. from Quebrada Honda.. somewhat projecting. all FM Brazil. Diagnosis. lying on a level between the small. WCAB 1705-8. the upper jaw extending a more little than superior.

no vocal sac. adult female. Hyla riveroi. skin of throat and chest smooth.. no tarsal ridge. hand. 9 mm. Dimensions.4 mm. when limbs are laid along the side. holotype. 6. 20 mm. traces of a skinfold across chest..3 mm. roof of skull not exostosed.4 mm. femur. heels overlap greatly. heel reaches middle of eye.. Body somewhat elongate.. knee and elbow meet snugly.. no dermal appendage on heel. Skin of head not co-ossified with skull.5 mm. . that of belly and lower surface of thigh uniformly and very finely granular. head length. Skin of upper parts highly smooth. than greatest width of head. 6. no distinct narrow glandular ridge encircling upper part of tympanum. CM 37433 (X 3). heel-to-toe.4 mm. head width. 14. 10. . in postaxillary region a little narrower — <^^V'''' Figure 29. tibia. Head and body. when hind legs are bent at right angles to body.— FROGS OF COLOMBIA —COCHRAN AND GOIN 285 a smaller but equally distinct rounded outer metatarsal tubercle.. when hind leg is adpressed. 6. A patagium extends from the back of upper arm to side of body.

Hyla microcephala. in a bush or herbaceous plant. At a distance the call of the male is a delicate. Calling individuals were invariably located above ground. The thighs are nearly immaculate.S. The holotype. but close at hand it has a more rasping quality. Hyla mathiassoni.: 286 U. with a headof 20 millimeters. pending Dr. 1959. Goin and Layne (1958. Remarks. 524. longitudinal rows resembling an hour-glass pattern. but since he graciously declined we take this opportunity of dedicating it to him. — CNHM 81779. Rivero now informs us that this frog is undoubtedly distinct from H. The following life history notes are quoted from Goin and Layne This is body length — (1958. forming an indistinct to distinct interocular bar. collected at Villavicencio. including thighs. We offered to let Dr. by Phillip Hershkovitz. Meta. faintly peppered with minute melanophores but with the pale yellow ground color being the most conspicuous element of the pattern. bia. NATIONAL MUSEUM BULLETIN —Dorsal 2 88 ground color ranging from a moderate brown on top of snout to pale brown on top of head and anterior portion of back to a yellowish on hind part of back and along sides. p. Each shank with three distinct but not well-defined transverse bars across their dorsal The upper surfaces of all limbs. but the dorsal surfaces of the shanks are generally distinctly crossed by about three blotches of dark brown. Entire ventral surface a very pale yellow. p. . Rivero describe this new species. 104) pointed out that their identification was tentative. misera and that it represents an undescribed species. Holotype. Dr. Juan Rivero's study of the species. There is generally a concentration of pigment on top of the head between the eyes. None of the males examined exceeds 18 millimeters in head-body length. Tiny brown spots concentrated into a rather indistinct transverse blotch above the scapular region and a rounded blotch above the sacral region. Colom- 1959. one of the smallest species of Hyla. p. surfaces. but it Variation. this species is comparable to Hyla ocularis of the southern United States coastal plain. 105) All specimens were collected in wet cutover land and partially flooded meadows. — exhibits a fair amount of variation in dorsal pattern. insect-like tinkle. The dorsal ground color is tan to light brown with a series of dark brown blotches on the back. In using the name "raisera" for this species. Color in alcohol. In voice and habits. is the largest known specimen. usually at a height of from two to four feet. new species Figure 30 —Stebbins and Hendrickson. The type series is rather uniform in structure. These blotches range from being asymmetrically arranged to being organized in two somewhat curving.

all MLS 76 AND GOIN (8). broadly rounded. rounded choanae. rounded patches. tongue three-fifths as wide as mouth opening. dorsolateral lymphatic — show through the dorsal skin and give a superficial appearance of broad dorsolateral stripes. the latter species do not have the conspicuous dorsolateral lymphatic sacs of mathiassoni. south of Villavicencio. lying close together between small. from Kio Ariari.— — —COCHRAN FROGS OF COLOMBIA CNUM Paratypes— 152215-35. 287 152105-7. Also. Diagnosis. its posterior border fused and very shallowly notched. from Florencia. holotype. Caqueta. USNM USNM 151962-3. Meta. maihiassoni at once from the related species in Colombia H. CJG 81775-8. . Meta. Snout short. USNM from Villavicencio. 152066-8. A small yellow frog of the microcephala group without a distinct dorsal pattern but with large. sacs that Description oj holotype. the upper jaw extending considerably beyond lower. riveroi. U-shaped when viewed from above. Hyla mathiassoni. 2337-9. —Vomerine teeth in two small. The absence of a distinct dorsal pattern serves to distinguish H. dark. microcephala and H. nostrils more lateral than superior. CNHM 81779 (x[3). somewhat pointed in profile. Figure 30.

when hind leg is adpressed. mm. NATIONAL MUSEUM BULLETIN 288 very slightly projecting.2 mm. no projecting rudiment of a pollex.288 U. the web on third finger reaching base of antepenultimate phalanx. A rather narrow glandular ridge encircling upper part of tympanum. loreal region flat and nearly vertical.6 mm. neither reaching to disk of third. which is most heavily (but pigment this the eye Posterior to centrated along the canthal region.. separated from eye by a distance nearly equal to its own diameter. somewhat lavender in color. Dimensions.S. . A moderately well-developed patagium extends from the back of upper arm to side of body. this loreal regions are slightly evidence of pigmentation on the limbs or venter. and greatly distended.7 mm. 10.. Skin of upper parts smooth. 8. heel reaches anterior margin of eye. There is no Color in alcohol. The constill conspicuously) not pale brown. continues as a narrow darkened strip that passes above the tympanum along the side to the sacral region where it fades out.1 mm. adult male. disk of third slightly more than covers the tympanic area.. that of belly and lower surface of thigh uniformly and very finely granular. hand. when limbs are laid along the side. finely wrinkled. heels overlap considerably. head width. a small distinct oval inner and a smaller but distinct rounded outer metatarsal tubercle. give a superficial appearance of pale lavender dorsolateral stripes extending from the region of shoulders to just above groin. and body. about one-third the diameter of eye. roof of skull not exostosed. separated from each other by an interval equal to their distance from eye.4 mm. Tympanum distinct. —Dorsally and ventrally specimen is a uniform pigmented with pale brownish yellow. 14. 6. no dermal appendage on heel.2 mm. Toes slightly more than one-half webbed.. femur. no ulnar ridge. heel-to-toe. Canthus rostralis moderately defined.4 —Head 6. tibia. no outer but a low inner tarsal ridge. Skin of head not co-ossified with skull. head length. in postaxillary region a little narrower than greatest width of head. Fingers webbed at base. the upper lip not flaring out appreciably below it. its diameter slightly greater than its distance from nostril. 6. Eye not large.. A pair of subdermal glandular(?) areas. which is relatively narrow and not as wide as distance between nostrils. 21. their distance from end of snout about onehalf that from eye. knee and elbow meet. a well-developed skinfold across chest. palpebral membrane not reticulate. fourth finger about equal to second. third and fifth toes subequal. when hind legs are bent at right angles to body. the web on fourth toe reaching base of penultimate phalanx.. interorbital distance greater than width of upper eyelid. disk of fourth toe just covering the tympanic area. vocal sac subgular. Body not elongate. skin of throat and chest smooth. but prominent due to black choroid coat of eye showing through translucent upper lid.

the judge whether the variation in pattern of impossible to phlebodes. 21. They give the following notes on the life history of the Villaviciencio frogs (Steb- Remarks. 1. there was little tendency for the eggs to separate at these zones.3 millimeters in the largest likewise. in diameter. The ova were about 1 mm. represents whether it species other than H. There is. in depth. that are characterized generally conspicuously by having a patagium. but pattern that in H. 1959. mathiassoni. —To our knowledge. p. In most individuals these sacs are very conspicuous. a variation in the conspicuousness of the dorsolateral subdermal lymph sacs. represents a Florencia population While most of the USNM The lymph sac showing through the skin of the dorsum surely represents the "very pale lavender stripe from snout to groin" as described by Stebbins and Hendrickson (1959. is due to from Florencia this specimen speciation. The zones of juncture between the outer capsules were made out with difficulty. in diameter with the lower quarter cream color and the remainder brown. and at 10:30 a. in bright daylight on November 20. and a pair of oval glandular areas on the chest. this be due to thicker or more heavily pigmented skin rather than to may less well-developed sacs. microcephala from Huila. ovoid mass. 526): A pair found in amplexus at Villavicencio was placed in a container in which the female deposited her eggs. female. but in others they are much less so. in fact.FROGS OF COLOMBIA Variation. 1950. Beyond this was a softer jelly that more or less merged with that of adjacent eggs but was set off sharply from the inner capsule. 14 of the 41 at hand show some degree of dorsal pigmentation.5 24. There were approximately 180 eggs. They were surrounded by a stiff jelly capsule. since less than a half dozen large ova remained in her oviducts. In four specimens this pigmentation simply amounts to tiny. specimens are without pigment on the back. It is simple geographic variation. AND 289 GOEST the greatest variation seems to be in hand range the smallest male to The specimens millimeters in —COCHRAN at in head-body length from 19. There are a dozen or more species in this group . pencil-like 152066. 525). the only published account pertaining to this species is that of Stebbins and Hendrickson (1959) in which four specimens from Villavicencio are compared with H. whether the incipient or. p.m. in width. size. measuring 37 mm. —Structurally. essentially her total complement. is LEUCOPHYLLATA group composed of small frogs. rather smoothsurfaced. thighs nearly or entirely without pigment. and about 9 mm. bins and Hendrickson. 21 mm. These were laid in a fairly compact.5 mm. This group patterned. Stebbins and Hendrickson heard the Villavicencio specimens call shortly after dark on November 18. in length. in a similar pigment is arranged to Florencia. from flecks scattered more or less over the back.

Ponte de Gentio on Rfo Al- 1829. A most a narrow broken H. less in Key a 1 .— 290 U. pi.— Shaw. on top of arms and legs. . ca . Iaynei rather distinct dark facial stripe region. 607. 1829.— Gunther 1858. 529. p. p. 671 (type locality. — Merrem. Upper Upper No 62. p. fig. A distinct pah such spots fused to form an of supraocular light spots or interocular bar. 31. 127. ebraccata definitive vomerine teeth. Hyla elegans Wied. tiny. — Dumeril p. p. 1803. 1901. 1789. — 243. 168. c1 . 1841. 1874b. Suriname) — — p. Gravenhorst. below eye not distinctly margined with white. dorsum uniform brownish yellow in life. more thickly placed. c1. 7. dorsum with bright gold spots H. p. Calamita leucophyllata. sometimes so thickly placed that the light areas between them form a network of fine lines H. pi. p. 1789. membranacea No dark brown stripe on side of face. H. or if present. NATIONAL MUSEUM BULLETIN 288 South America. 1868. b2. 11. 1820. Bonnaterre. 1799. b 1 . Schneider. H. dorsolateral light marks restricted to a row of light dots or at stripe c2. fig. Definitive c1 . than eight of these are known from Colombian Species to in the leucophyllata Group Dorsal surface dark with a pattern of light gray or silvery white light marks. 156. sarayacuensis along the sides of body Dorsal surface plain or with a distinct pattern but without a pattern of silvery white or gray marks. 4. 1789. Rana variegata Bonnaterre. oliveae in life Dorsal pattern conspicuous. p. Dorsal pattern of thickly scattered dark spots.— Steindachner. 1820.— Cope. Gmelin.H. 277. Surinam). 4. 1055. Hyla leucophyllata. 1864. p. c1 . p. 1824. 222. . 173. fig. Hyla leucophyllata (Beireis) Figure 31 1783. 1802. 45. 182. and not b 1 . fig. rather distinct dorsolateral light stripes extending for one-third to one-half the distance to groin a2 . Rana leucophylla. 1. unknown). p. cobaca). 2. reticulata in a reticulate light pattern Dorsal pattern of dark spots restricted to one or two rows down to middle of back and others. 1 (type locality. Calamita leucophyllatus. but rest of top of head not light. c2. 85. lip well developed. 17. 1824. H. (in life) vomerine teeth. p. A brown on side of head and loreal and tympanum to taper out along the side. Rana leucophyllata Beireis. pi. 1925. rossalleni light triangular area covering top of head. p. and no Colombia. p.S. p. 4 (type locality. p. 8 (type locality. extending posteriorly through eye c2 . pi. 1871b. 112. Hyla frontalis Daudin. leucophyllata lip below eye distinctly margined with white H. p. inconspicuous. Dorsal pattern obsolete or (in life) made up of bright gold spots. . p. 1789. and Bibron. Ill. 1802. 1799. 1870.

Figure 31. 31. 1926. 22. p. 44. 1892. 1870. 1838. 9. 6. pp. an adult female from Florencia. 1937. 3a.— Mertens. p. figs. pi. 3.— Gorham. Hyla leucophyllata.— Cott. Shreve. light locality. p. Cochran. lip below the eye distinguishes ebraccata. p. 1930. 1950. 1963. p. 1935. p. 115. sarayacuensis and H. p. 55. fig. p. —COCHRAN AND GOLN 291 387— Boettger. p. Fitzinger. 1923. 31. 38. 105. . Description. Brazil). — — Hypsiboas leucophyllatus.— Baumann.— Mertens. 1958. 292. MLS 35b (X 2). pi. Cope. The absence of a distinct white margin on the upper it from H. 227.— Myers. white daub that covers the dorsal surface of the shank serve at once to separate this species from both H. p. 1946. 1882a. — — — 1838. 10.— FROGS OF COLOMBIA 122— Boulenger. Hyla triangulum Gunther.— De Witte. p. fig.— Nieden. 72. p. p. p. Diagnosis. The conspicuous light marks on the dorsum that surround a dark rectangular or triangular blotch and the large.— Kreig and Forster. Caqueta. 13. p. figs. 1930. 259. Goin and Layne. p. 489. p. 1926. Dendroscophus frontalis. 187. 4 (type Hyla leucophyllata triangulum. 1912. 1869. p. 162. 215. Tschudi. — MLS 35b. fig. pi. 1160— MirandaRibeiro. — —A member of the leucophyllata group with conspicuous markings dorsally. a-d. rossalleni. 1955. 73. 163. p. 9. 1843. 1843.

S.. no vocal sac. heel reaches anterior margin of eye. separated from each other by an interval equal to about two-thirds their distance from eye. 10 mm. head width. 21.. when hind leg is adpressed. just reaching to disk of third which just covers the tympanic area. Fingers webbed at base. Color in alcohol. not particularly prominent. which is slightly greater than distance between nostrils. palpebral membrane not reticulate. third and fifth toes subequal. that of belly and lower surface of thigh uniformly and very finely granular. when limbs are laid along the side. 26. knee and elbow overlap considerably. Can thus rostralis well defined. their distance from end of snout about one-third that from eye. Tympanum distinct. head length. rounded choanae. its posterior border very slightly free and unnotched. broadly cordiform.6 mm. Snout short. nostrils more lateral than superior. Dorsally a rich chocolate brown with a distinct pattern of faded creamy white. the upper jaw extending but very slightly beyond lower. heel-to-toe. 13.. heels overlap greatly.. Head and body.6 mm. Skin of head not co-ossified with skull.4 mm. femur. disk of fourth toe covering about two-thirds the tympanic area. then passing downward to terminate above arm. includes all of upper lid and continues posteriorly from corner of eye above the tympanum to a point just — — . web extending to distal third of antepenultimate phalanx of third finger.8 mm. A patagium extends from the back of the upper arm to the side of the body. the upper lip not flaring out below it. no projecting rudiment of a pollex. This dorsal pattern consists of a sharply defined pale white area that extends from the nostrils along the canthus to anterior corner of eye. no tarsal ridge. 16. its diameter equal to its distance from nostril... skin of throat and chest smooth. rounded when viewed from above. no ulnar ridge. roof of skull not exostosed. Skin of upper parts smooth. Toes more than one-half webbed. Eye moderate. slightly projecting. NATIONAL MUSEUM BULLETIN 288 Colombia. fourth finger considerably longer than second.6 mm. a welldeveloped skinfold across chest. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Vomerine teeth in two tiny rounded patches lying between the small.292 U. in postaxillary region a little narrower than greatest width of head. the web on fourth toe reaching base of penultimate phalanx. about one-third the diameter of eye. hand. when hind legs are bent at right angles to body. truncate in profile. tibia.1 mm. adult female. no dermal appendage on heel. tongue one-half as wide as mouth opening. Dimensions. separated from eye by a distance about equal to its own diameter. interorbital distance slightly greater than width of upper eyelid. Body not elongate. loreal region flat and very nearly vertical. a glandular ridge encircling upper part of tympanum. 10. 9.

Outside of Colombia. and. Along the sides the dark dorsal ground color merges gradually with the cadaverish gray ventral color. one on the for it has two rather large distinct daubs on each shank — 337-262—70 20 . and the apex extends posteriorly about halfway to the sacral hump. which is just west of Florencia. Furthermore. of course. — Remarks. Specimens from Florencia have the rectangular dark spot reduced to an elongate. 31). extending from knee to tip of heel. Pigment on thigh is limited to a trace on the distal portion of is dorsal surface. It is impossible. shanks (including both knee and heel) and fore arm are white. bifurca. Since one can find specimens similar to bifurca in eastern Ecuador together with sarayacuensis and leucophyllata. A conspicuous. but not quite. comes in contact with the posterior end of the dorsolateral stripes in the region of the sacral hump. individuals that match Andersson's description and figure of H. Specimens from Leticia have the pattern that usually is considered rather typical for the species top of head and dorsum silvery white with a dark rectangular figure extending from between the eyes to region of the sacral hump and a large. The type of Hyla frontalis is somewhat intermediate in pattern between bifurca and leucophyllata. is between the eyes. the marking of the shank is intermediate. In this region one can find fairly typical examples of H. on the present state of our knowledge. leucophyllata. sarayacuensis at Sarayacu. but specimens from the eastern slopes of the Andes in Ecuador and Peru will have to be studied in more detail than previously before a real understanding can be reached. body. it differs in that there is no connection between the white above the vent and the white in the dorsolateral lines and in that the white daub is absent from the dorsal surface of the shank and is restricted to a tiny spot on each heel. Specimens from southeastern Brazil seem to be essentially like those from eastern Colombia. triangular spot that extends from between the eyes and has its apex reaching about the middle of the back (see Fig. While bifurca superficially resembles leucophyllata. light-colored area is present on dorsal surface of each shank. the base of the triangle. it seems probable that bifurca is a valid species. Two specimens (MLS 225-6) from Ceilan. silvery white daub completely covering the dorsal surface of the shank. an adequate discussion of the variation in Hyla leucophyllata. In it the white spot above the vent is quite enlarged and nearly. the problem of geographic distribution becomes even more complex.FROGS OF COLOMBIA —COCHRAN AND GOIN 293 above the vent. An elongate triangular area of the dark ground color present on the back. have the dorsal to write — dark spot completely obliterated so that the entire dorsal surfaces of head. topotypes of H.

1945. is usually constricted in the middle (and thus dumb-bell shaped) rather than gradually tapering posteriorly as in leucophyllata. JAP 2697. p. leucophyllata. . its diameter equal to its distance from — . west of Puerto Napo. rounded series. from Rio Manso.S. Hyla ebraccata is most similar to H. MLS 292... slightly projecting. its posterior border slightly free and shallowly notched. JAP 2638. 35b. 2745-6.— Taylor. A member of the leucophyllata group with a white Diagnosis. MCZ 4477. Nicaragua). 1881. Caqueta: MLS Florencia. Antioquia: Rfo Sucio. CNHM MLS 8526 8527 (2). lying between the small.— Taylor and Smith. 8525 (3). MCZ 16273-4. — Hyla ebraccata Cope. MCZ 17598. WCAB MCZ 152283. broadly rounded. 225-6.— Taylor. rounded choanae. tongue two-thirds as wide as mouth opening. 44. 30. p. 69 (type locality. Ceilan. — — — 1960c. Napo-Pastaza: Two km. the top of the shank is not completely covered by a daub as it is in leucophyllata. Amazonas. 1942. Bonita. No exact BRAZIL: USNM locality: Rfo Putumayo. Cundinamarca: Mountains of Medina. D. frontalis) Hyla ebraccata Cope Plate 37a-c 1874. loreal region nearly flat and nearly vertical. p. 486. Can thus rostralis rather well defined. 465 m. from which it may be distinguished by having a conspicuous white margin on the upper lip just below the eye and the fact that the dorsal dark blotch. USNM 152064-5. 2824 (7) Manquinhos. — p. NATIONAL MUSEUM BULLETIN 2 88 — proximal end and one on the distal end and a tiny daub on the heel. Eye moderate. Acre. if present. northeast of Tena. one km. Colombia. Thus.1960. south bank Rio Napo. 1942. p. their distance from end of snout about one-half that from eye.. 885 Langebartel and Smith 1959. northwest of Villa vicencio. separated from each other by an interval nearly equal to their distance from eye. no data. Brocchi. not prominent. p. CJG 2369. MCZ 2030. Duellman. Snout rounded when viewed from above. F. 40. 82014. Specimens Examined COLOMBIA Amazon as: UF Leticia. 1262. and it is not dark as it is in bijurca. 27. C<5rdoba. SURINAME: MHNP 4868 (type of H . Hyla leucophyllata. west of Florencia. 1874a. Fouquette. 1952. nostrils more lateral than superior and as anterior as lateral. Vomerine teeth in two widely spaced. Pernambuco. margin on the upper lip just below the eye. p. 587. the upper jaw extending but little beyond lower. (2). 294 U. p. slightly rounded in profile. the upper lip not flaring out below it. PERU: Ambyiaca River. Description. ECUADOR: JAP 2726.

Skin of head not co-ossified with skull.4 mm. when hind legs are bent at right angles to body. Head and body. no idnar ridge. a low. The thighs of hind limbs are without pattern. when limbs are laid along the side. interorbital distance than width of upper eyelid. 10. 20. as is characteristic of frogs of this group. dorsum between the dorsolateral stripes thickly spotted with tiny white spots. inconspicuous fold encircles upper part of tympanum. no vocal sac. knee and elbow overlap considerably.6 mm. rounded glandular areas in the pectoral region. posed pattern of silvery white.. A well-developed patagium extends from back of upper arm to side of body. just reaching to disk of third which covers half again the tympanic area. a horizontal white streak occurring on each upper jaw and extending from eye to edge of lip. tibia. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Tympanum very distinct. Body not elongate.9 mm.. no projecting rudiment of a poll ex. All ventral surfaces are immaculate.FROGS OF COLOMBIA —COCHRAN AND GOIN 295 membrane not reticulate. heel reaches tip of snout. about one-half the diameter of eye. On the anterior limbs there are several small white spots with a larger one just proximal to the wrist. femur. Toes almost completely webbed. Dorsal ground color bluish gray with a superimColor in alcohol. heels overlap moderately.. but there is a small white spot on each knee.8 mm. and in each notch there is a small white spot.. 28. traces of a skinfold across chest. 36.7 mm. a heart-shaped spot on dorsum extending from region of the sacral hump to above the vent. and on the dorsal surface of each shank is an elongate white daub with a distinct notch on the inner side of each daub.7 mm. roof of skull not exostosed. two distinct. in postaxillary region about equal to the greatest width of head. 6. heel-to-toe. third and fifth toes subequal. 19.3 mm. fourth finger only slightly longer than second. Skin of upper parts smooth. Fingers webbed. palpebral slightly greater — — . Dimensions. nostril. no tarsal ridge. adult female. 11. separated from eye by a distance nearly equal to its own diameter. disk of fourth toe slightly more than covering the tympanic area. the web on fourth toe reaching base of penultimate phalanx. The lateral surface of the left foot is margined in white. the web on the third finger reaching the base of penultimate phalanx on both sides. that of belly and lower surface of thigh uniformly granular. The silvery white pattern is comprised as follows: a triangular area on top of head with the posterior corners of the triangle each continuous with a broad dorsolateral stripe that passes above eye and tympanum and terminates about the level of the sacral hump. which is about equal the distance between nostrils. skin of throat and chest smooth. no dermal appendage on heel. when hind leg is adpressed. head length. head width... hand.

p. In pattern there is great variation. 1957. and in lacking large. distinct. Diagnosis. Two fresh females from the Rio Manso. the choanae small. narrows as it passes posteriorly. while the four largest males from the same series (CJG 2368-82) measure 26. 215 (type locality. 1958. Hyla leucophyllata. 1955. In the female from the Rio Sucio. —A brightly colored frog of the leucophyllata group with a dorsal pattern of light From Hyla marks on a dark background. In all other specimens from the Rio Manso three basic types of pattern are noted. CJG 2368-82. leucophyllata it differs in lacking a large. leucophyllata sarayacuensis Shreve. NATIONAL MUSEUM BULLETIN 288 —As in Hyla leucophyllata. measure 36. and 29. scattered brown marks on the back and a brown interocular bar. Amazonas. — Goin and Layne. Hyla sarayacuensis Shreve Figure 32 1935.S. in the size of breeding individuals of the Specimens Examined COLOMBIA Antioquia: Rio Sucio. C6rdoba: Rio Manso. Cordoba. rossalleni. there is a marked difference two sexes. — Cochran. From H. which is probably its nearest relative. somewhat oblong. Antioquia.3 millimeters. the top of the head and dorsum are white with about a half-dozen small. The female described and figured above is the most gaudy of the lot. thus forming a sort of hour-glass figure on the back. 106. CNHM 82014. then widens and terminates above the sacral hump. — Goin. Sarayacu. p. white supraocular spots. 28. H. Variation. Vomerine teeth absent. Hyla 1955. In the third type the general tone of the dorsal ground color is light brown with a dark brown dorsal figure that begins between the eyes. In all of the specimens the white mark on the upper j aw between the lip and the eye is present. 1957.296 U. rounded.5. UF 8521-1. Ecuador). . In another the entire top of the head and dorsum and the upper surfaces of the shanks are tan. — Description. p.3. in having a triangular pale area on top of the head. bright. 60. an adult male from Leticia. 27. Hyla sarayacuensis. tongue three-fourths as wide as mouth opening. its posterior border slightly free and shallowly notched. sarayacuensis differs in having the dorsolateral stripes complete.8 and 38 millimeters in head and body length. p. Colombia. rectangular or triangular spot on the back that is completely surrounded with white and in lacking the large white daub on each leg that nearly covers the dorsal surface of the shank. 115. Snout short. In one type the entire top of the head and dorsum and the tops of the skanks are silvery white.

which is relatively wide and slightly greater than distance between nostrils. fourth finger considerably longer than second. Eye large. its diameter equal to lKjtimes its distance from nostril.— FROGS OF COLOMBIA —COCHRAN AND GOIN 297 rather rounded when viewed from above. interorbital distance slightly greater than width of upper eyelid. separated from eye by a distance nearly equal to one-third its own diameter. no ulnar ridge. Canthus rostralis slightly defined. separated from each other by an interval equal to about three-fourths their distance from eye. reaching to base of disk of third which snugly covers the tympanic area. slightly projecting. Fingers webbed to proximal end of penultimate phalanx of third finger and to middle of penultimate phalanx of fourth. no projecting rudiment of a pollex. truncate in profile. the web on fourth toe reaching distal end of antepenultimate phalanx. Tympanum distinct. the upper lip not flaring out below it.. Figure 32. . their distance from end of snout about one-third that from eye. UF 8521-1 (X 2). the upper jaw extending but very slightly beyond lower. nostrils more lateral than superior. disk of fourth toe covering about three-fourths the tympanic area. an oval inner and a smaller . about one-third the diameter of eye. palpebral membrane not reticulate. loreal region somewhat rounded and nearly vertical. very prominent. third and fifth toes subequal. Toes slightly more than one-half webbed. Hyla sarayacuensis.

the dorsal ground color pattern of gray (silver in is gaudy frog. skin of throat and chest smooth except for the pair of pectoral glandular patches which are characteristic of this species. and the posterior margin of the lower jaw.6 mm. tibia. 25. a rather narrow glandular ridge encircles the upper part of tympanum. 9... : extends posteriorly for nearly half the length of the body.. no dermal appendage on heel. and a crossband on the middle of the crus and the upper segment of the foot. it covers the dorsal surface of the crus and the lateral third of the foot and extends out onto the two lateral toes. when hind legs are bent at right angles to body. a light spot on each light dots elbow and wrist. It also covers the tympanum and lower arms. On the hind limb it is absent from the thigh except for a very narrow streak on the distal end of the dorsal surface. — Color in alcohol. Skin of head not co-ossified with skull. The dorsal ground color extends over the entire face.298 U. heel reaches beyond tip of snout. 20. There is is imposed a a very sharp line of demarcation between the dorsal ground color and the completely pigmentless ventral surfaces.6 mm. roof of skull not exostosed. subgular pouch with longitudinal folds. in postaxillary region distinctly but not appreciably narrower than greatest width of head. a well-developed skinfold across chest. A patagium extends from back of upper arm to side of body. hand.S. NATIONAL MUSEUM BULLETIN 288 rounded outer metatarsal tubercle. Body not elongate. The head-body length of the larger females ranges from 29 to 34 millimeters while that of the smaller males ranges from 23 to 25 millimeters. head width.9 mm.5 mm. that of belly and lower surface of thigh uniformly and very finely granular. vocal sac.1 mm. a light spot on each knee. Dimensions. 7. upper lips. Head and body. 14. On the back the posterior edge of the it extends laterally to a line drawn from to the vent. On this dorsal ground color the following light pattern is imposed a triangular area on top of the head between the eyes and nostrils.. a postocular light stripe which dorsal surface of the upper half of the hand. scattered and streaks on the dorsal surface. The most conspicuous difference that can be noted in the Colombian specimens is that of size. the top of the lateral and extends out onto the lateral two fingers. Skin of upper parts smooth.. femur. no tarsal ridge. —A brightly colored. heels overlap greatly. 9.4 mm. head length. knee and elbow overlap considerably. when limbs are laid along the side. — Variation. In life the ground color of the dorsum in this species is brownish. 13 mm. However. heel-to-toe. when hind leg is adpressed. top of head. and ankle joint and on the base of the fifth toe. In preservative medium brown on which life) markings. A triangular patch .. heel. while the upper parts are bright salmon pink.

south-southeast of Puyo. alleni. white. rossalleni. sarayacuensis never seems to have the large. p. Rio Puyo at Puyo. generally a foot or more above the ground. p. There is a definitive spot just above the vent in some specimens. and in lacking the bright. of a dark frog with light markings. silvery white supraocular spots. 19729 (type of sarayacuensis). In the two that do not show this condition. 1959. Goin and Layne. Furthermore. pre- — occupied). In the latter the creamy white markings are so extensive on the dorsum that the impression is given of a light frog with dark markings. In all except two of the Leticia specimens there are two small spots in the sacral region. sarayacuensis were collected on vegetation in a wet meadow. While about the same size as leucophyllata it differs in its more marked sexual dimorphism. All of the Leticia specimens of H. triangular spot on top of the head and snout. In H. 60 (type locality. H. rossalleni in being much larger in size. This light patch is missing in one specimen. p. silvery daub on the dorsal surface of the shank that is so characteristic of leucophyllata. each pair on the large leaves of an aroid (Araceae) plant. in having the dorsolateral stripes more or less complete. Hyla alleni Goin. . 2979. no additional specimens were seen. that is. leucophyllata and H. 19730 (paratype of sarayacuensis) EPN 2975-6. 106. 1959b. Three pairs were found in amplexus. 1958. — COLOMBIA Specimens Examined r Amazonas: Leticia. and an aggregation of creamy white flecks in the same position in others. JAP 2055. Amazonas. JAP 2009. ECUADOR: Napo-Pastaza: Sarayacu. Remarks. MCZ MCZ MCZ . Hyla rossalleni Goin Figure 33 1957. UF 8521 (9). in lacking definitive vomerine teeth.— FROGS OF COLOMBIA —COCHRAN AND GOIN 299 on top of the head from the eyes to the nostrils is a creamy white. 1963. Leticia. 21. Although obviously related to both H. A band of creamy white crosses the shank. A line of similar color extends from behind the eye to about midbody. sarayacuensis the opposite impression is gained. this species is just as obviously distinct from both. They were all taken on a single night. Hyla rossalleni Goin. lower Rio Apaporis. and in pattern. sarayacuensis differs from H. and patches of similar color occur on the heel and elbow in all the individuals. 340 (substitute name for H. Gorham. the spots are fused into a dumb-bell shaped figure in one and are not visible at all in the other. Vatjpes: Gino-Goj'6. Colombia). in having a white. and though the same and similar-appearing habitats were worked on other occasions and under seemingly similar conditions. p. 6 km. passing above the tympanum and axilla. 28047.

Eyes large and protuberant. by the presence of the supraocular spots. its diameter about two-thirds the width of mouth. Hyla rossalleni. dorsolateral light stripe broken spots. sarayacuensis. snout nearly rounded as seen from above. From H. tongue nearly round. Tympanum distinct but small.— 300 U. diameter of eye greater than the distance from eye to nostril. Head broad. from Leticia. and well separated. lombia. CoDescription. leucophyllata it differs by lacking vomerine teeth and by lacking the large white daub which covers the dorsal surface of the cms. distinguished from H. rounded. Amazonas. not more than one-third the diameter of eye. NATIONAL MUSEUM BULLETIN 2 88 — Diagnosis. UF 8501 (holotype). A small Hyla of the leucophyllata group with conspicuous supraocular light spots or interocular bar and without a triangular light area H. nostril very near tip of snout. its nearest by having the up into more or less elongate light and by its smaller size. not notched behind and but slightly free behind. on top may be of the head. its width greater than distance from snout to occiput. UF 8501 (X 2). upper jaw very slightly projecting. . choanae moderate in size. — Figure 33. rossalleni relative.S. Vomerine teeth absent. interorbital space about equal to diameter of individual eye.

its fellow from the opposite side. The areas are separated on the median line by about two millimeters. 7. . toes 4-3-5-2-1 in order of decreasing length. 22. Well-develdisk of second finger about equal third finger of left hand partially of terminal phalanx). Dimensions. above each eye.8 mm. A moderately developed patagium extends from back of upper arm to side of body. A few — . 38. underside of chin covered by loosely folded.. Venter and underside of thighs rugose. diameter of tympanum. upper eyelids. fully webbed except the fourth toe which has the web extending to base of penultimate phalanx. There is a pair of glandular areas in the skin of the pectoral region. Color in life. There is. The entire ventral surface is immaculate except for a few flecks of dark pigment on each side of the throat near the posterior portion of the lower jaw.3 mm. creamy patch that extends from the free margin of the upper lid down onto the interocular region where it approaches. When captured. 8. Head and body. The thighs were salmon pink below and somewhat dusky dorsally. arched row of light flecks that extends from above the tympanum on each side to above the axilla where it becomes obsolescent. punctulations of darker brown on the back between the eyes and the shoulder region and a similar patch of darker punctulations above the sacral region. knees and elbows in contact when limbs are pressed along the side. Darker pigment is also in evidence on the elbows and in bands across the shank. — — . Heels slightly overlapping when legs are flexed with femora held at right angles to the body.3 mm. The bars above the eyes and the row of light flecks on each side behind the head were creamy white.third its length web extends distally to base of penultimate slender. Similar creamy white patches make up a short. head width. (disk of tymand dorsum smooth.. 3 mm. but does not quite meet. heel extending to anterior margin of eye when leg is pressed along body. 1 mm. external vocal pouch which is very finely rugose.. top of head. diameter of eye. this specimen was predominately brown above and whitish below. Ground color above tannish brown.FROGS OF COLOMBIA — COCHRAN distance from tympanum panum.7 mm. fingers 3-4-2-1 in order of decreasing length with fingers 4 and 2 being nearly subequal. Each area is about 2% millimeters in diameter and lies just posterior to the insertion of the arm. third finger fully but a slender margin of phalanx. tip of snout to posterior margin of tympanum. hind leg (vent to tip of longest toe). Toes slender. Fingers not particularly webbed for only about one.. a conspicuous. hind leg (vent to heel). 22. The dorsal surface of the thigh has on it some tannish pigment similar to the dorsal ground color but is nevertheless paler than the dorsum and is without a distinct pattern. scattered Color in alcohol.5 mm. oped disks on all to diameter of removed to permit view 301 to eye slightly less than diameter of and fingers tympanum AND GOIN toes.

p. the upper lip not flaring out strongly below it. Brazil. about one-third the diameter of eye. 1952. slightly rounded in profile. Tympanum very distinct. p. and sometimes they are fused in the midline to form an interocular bar. their distance snout about one-half that from eye. slightly projecting. Lutz and Kloss. UF WCAB 8501 (holotype). p. from which on the back and in having dark loreal regions. 106. 1414. and a Hyla membranacea it differs is in lacking gold spots different voice.S. a uniform brown dorsum and a dark loreal and postocular band that progressively diminishes along the side as it approaches the groin. — UF 8523. Canthus rostralis slightly defined. separated from eye by a distance nearly equal to its own diameter. The only noteworthy variation we have seen in specimens is in the supraocular marks. Although no additional material has been taken in Colombia since the type series was collected. palpebral membrane not reticulate. broadly cordiform. the web on third finger reaching base of penultimate phalanx. 651. Bokermann has taken some specimens in Acre. loreal region nearly flat and very nearly vertical. 77 (type locality. 1958. Rio Pastaza). the upper jaw extending slightly beyond lower. these have faded — Remarks. Eye relatively large for a frog this size. Description. rounded series. which is relatively wide and slightly greater than distance between nostrils. p. NATIONAL MUSEUM BULLETIN 2 88 tiny flecks of a similar nature were present along the shank although somewhat in preservative. its diameter slightly exceeding its distance from nostril. Sometimes these are discrete spots. perhaps most similar to H. separated from each other by an interval equal to about their distance from eye. tongue one-half as wide as mouth opening. lying close together between the small. Specimens Examined COLOMBIA Amazon as: BRAZIL: Leticia. Acre. very broadly triangular when viewed from above. ine teeth in more lateral than superior. oliveae. Goin and Layne. — A member of the leucophyllata group characterized by Diagnosis. one on top of each eye. Hyla membranacea Andersson. fourth finger but slightly nostrils from end of . Colombia. — — 1963. Hyla membranacea Andersson Plate 37d-f 1945. 22. Snout short. from Leticia. interorbital distance about equal width of upper eyelid. Ecuador. Fingers webbed. a dark postocular stripe. Vomertwo short. a more triangular head. and was kind enough to send us one for examination. Amazonas.— 302 U. Gorham. its posterior border free and with a trace of notch. rounded choanae. 8502 (paratype).

— are remarkably uniform. There are several little darker spots on the back but these are so inconspicuous and scattered that in no sense can the specimen be said to of thigh — — have a spotted dorsum.. Body not elongate.4 mm. The four specimens in the University of Florida series Variation.6 mm. . a rather narrow glandular ridge passing above upper part of tympanum. —In life all thigh is the specimens were an unmarked brownish yellow on the back with a blackish subocular and postocular stripe that extended to the groin as a narrow dusky stripe. when hind legs are bent at right angles to body.7 mm. The distended vocal sacs were yellowish white. and this dark brown coloration passes posteriorly through the tympanum and above the shoulder. 9. heel-to-toe.4 mm. then continues posteriorly as a progressively fading stripe nearly to the groin. a distinct oval inner and a smaller indistinct rounded outer metatarsal tubercle... there are no noteworthy differences between the individuals. no dermal appendage on heel. So far as pattern is in size from concerned..3 millimeters. rounded glandular areas in pectoral region. A patagium extends from the back of upper arm to side of body. when hind leg is adpressed. 13. a pair of well-developed.. toes subequal.—COCHRAN FROGS OF COLOMBIA AND GOIN 303 longer than second. Color in life. They are all males and range 22. that of belly and lower surface no tarsal ridge. heel reaches anterior margin of eye. in postaxillary region a little narrower than greatest width of head. Skin of upper parts smooth. skin of throat and chest finely wrinkled. Except for the margin of the lower lips. 26. 9. Head and body. heels overlap. Dimensions. Skin of head not co-ossified with skull. 11. as is the upper The unpigmented except for a narrow band of pigment along its dorsal surface connecting the pigment of the dorsum with the pigment on the dorsal surface of the shank. femur. no projecting rudiment of a pollex. adult male. when limbs are laid along the side.4 mm. The upper Up and loreal region are dark brown. Color in alcohol. the web on fourth toe reaching base of penultimate phalanx. The dorsal surface of both the upper and lower arm is heavily pigmented.. yellowish gray below. Pale brown above. no ulnar ridge. and the palms and soles were salmon pink.3 mm. hand.7 to 26. tibia. roof of skull not exostosed. just reaching to disk of third which slightly more than covers the tympanic area. 19 mm. immaculate. a well-developed skinfold across chest. knee and elbow touch snugly. disk of fourth toe snugly covering the tympanic area. uniformly granular. 9. Toes more than one-half webbed. head width. vocal sac apparent only as wrinkled skin in gular region. The ventral surfaces of the body. the ventral surfaces are surface of the shank and foot. head length. third and fifth. except for glandular areas mentioned above. the thighs.

indistinct series. ere-e-e-e-e-e-p and very similar to that of Remarks. and in these the same facial mask can be seen exactly as it is in the Leticia specimens. Specimens Examined COLOMBIA Amazon as: ECUADOR: Leticia.304 U. rounded choanae. A Hyla of the leucophyllata group with well-developed patagia. in lacking dark lores and a dark postocular stripe. Amazonas. The brown dorsum with life seems to be unique. (4 specimens. no significant differences between the two series can be discerned. 8524 (2). tongue three-fourths as wide as mouth opening. size of tympanum. Two specimens in the type series have not females) or whether it cannot now be faded as much as the others.—Vomerine teeth in two short. new . and in voice. we were and compare from Leticia in the able to examine the four specimens in the type series them directly with the four fresh specimens University of Florida collection. UF Rfo Pastaza. collected by E. The only appreciable difference between the two series is that of size. This new species perhaps is most similar to H. NATIONAL MUSEUM BULLETIN 288 — Life history notes. Alf G. In structural characters such as teeth. Whether this variation is sexual difference (the Leticia specimens are all due entirely to males. The voice is a loud delibrate cre-e-e-e-e-e-p. Snout .7 millimeters while that of the specimens from Leticia ranges from 22. types of H. membranacea. the types all may be due in part to geographic variation determined. its posterior border free and very shallowly notched. leucophyllata. 8522-3. Diagnosis.' H.S. The head-body length of the type series ranges from 35. lying close together between the small. Two were collected from vegetation growing stream that bordered the meadow.3 millimeters. from which it differs in having the bright gold spots on the dorsum. Layne on February 23. —Thanks to the kindness of Dr. very broadly cordi- form. shape of head. 1956. from Leticia.4 to 36. Colombia. and amount of webbing. Johnels. species Figure 34 — — Holotype. in having a more gold spots in rounded snout.7 to 26. Description of holotype. UF 8555. Ross Allen and James N. pectoral glands. The other two were collected while they were calling in the meadow of the four individuals taken in water along the margin of a proper. membranacea) NR Hyla oliveae. tongue. and vomerine teeth. The University of Florida specimens were collected while they were calling along the edge of a wet meadow.

FROGS OF COLOMBIA short. 3. a distinct oval inner and a smaller indistinct rounded outer metatarsal tubercle. knee and elbow meet snugly. not strongly little rounded and nearly vertical. —COCHRAN rounded when viewed from above. holotype. interorbital distance about equal to width of upper eyelid. when hind legs are bent at right angles to body. UF 8555 (X 2). no ulnar ridge. separated from eye by a distance defined. Fingers webbed at base. no tarsal ridge. disk of fourth toe very nearly covering the typanic area. heels overlap. A patagium extends from the back of upper most to anterior Figure 34. Eye large and prominent for a frog this size. when limbs are laid along the side. when hind leg is adpressed. separated from each other of snout by an about one-half interval equal to about their distance from eye. just reaching to disk of third which snugly covers the tympanic area. their distance from end that from eye. third and fifth toes subequal. no dermal appendage on nearly equal to heel. . Hyla oliveae. Body not elongate. the upper lip not flaring out below it. its long diameter definitely greater than its distance from nostril palpebral membrane not reticulate. the web on fourth toe reaching base of penultimate phalanx. Toes more than one-half webbed. heel reaches al- margin of eye. about one-third the diameter of eye. which is relatively wide and slightly greater than distance between nostrils. the upper jaw extending beyond lower. the web extending to or onto the bases of penultimate phalanges of digits 2. no projecting rudiment of a pollex. AND GOIN 305 slightly pointed in profile. Canthus rostralis rounded. nostrils as lateral as superior. and 4. Tympanum very distinct. fourth finger considerably longer than second.— . in postaxillary region a little narrower than greatest width of head. slightly projecting. loreal region a its own diameter.

. We are happy to name this species for Olive Bown Goin. a pair of rounded glandular areas in the pectoral region. 1923. The specimen is uniform tan on the top of the head and dorsum.7 mm. This specimen was collected in open. These golden spots have all but disappeared in preservative. 306 arm to side of body. — Diagnosis. hands. The thighs are immaculate except for a thin row of tan flecks along the dorsal surface.7 mm.5 mm. 11 mm.7 mm. Lutz and Kloss.NATIONAL MUSEUM BULLETIN 288 U.. 7a. 61 (type locality. —When collected this specimen was a rich brown above.S. Hyla — 1963.— Nieden. but he refrained from naming it in the hope that additional material would become available. The upper surfaces of lower arms. 1923. 1952. p. Hyla reticulata Jiminez de la Espada Plate 37g-i 1870. figs. p.. 1882a. dotted with discrete. The ventral surface Coloration in otherwise immaculate. p. 8. p. head length. Skin of head not coossified with skull. 1957). 52. heel-to-toe. vocal sac apparent only as a loose gular fold. femur. No one has yet been successful in collecting more specimens and we cannot see that anything is to be gained by withholding the description any longer. and the tops of the feet are densely mottled with tan. p. skin of throat and chest smooth except for glandular areas mentioned above. hand. 16. 95 (type locality.269— Myers. 386. — — Nieden. tibia. wet pasture lands. roof of skull not exostosed. head — width. Rio Napo. p. Pebas. adult male. Ecuador).5 . is life. Gorham. Remarks. reticulata Jiminez de la Espada. Dimensions. Ecuador). p. 23. — Color in alcohol. The junior author suspected that this specimen was a novelty at the time he described other species from the same collection (Goin. that of belly and lower surface of thigh uniformly and very finely granular. 7. 1875. being comparable to the noise produced by rubbing the teeth of a comb. p. The upper lip and loreal region are brownish rather than tan. 8. Skin of upper parts highly smooth.— Boulenger.. 1886.. 24 mm. Hyla favosa Cope. shanks. A small frog of the leucophyllata group with a dorsal pattern of thickly scattered dark spots that sometimes are so closely . a rather narrow but well-developed glandular ridge encircling upper part of tympanum. 269. Head and body. there are a few flecks of tan along the margin of the lower jaw. 6. small golden spots. in recognition — of the help she has given us on this manuscript. 12. 1942. mm. a well-developed skinfold across chest. 652. Its call was loud and Pseudacris-like.

Skin of upper parts smooth. separated from eye by a distance nearly equal to its own diameter. Eye moderate. no projecting rudiment of a pollex. the upper jaw extending but very slightly beyond lower. skin of throat and chest smooth. rounded when viewed from above. Description. that of belly and lower surface of thigh uniformly and finely granular. Caqueta.2 Color in alcohol. about one-third the diameter of eye. tongue three-fourths as wide as mouth opening. rounded choanae. hand.. their distance from end of snout about one-half that from eye. west of Florencia. reaching to proximal third of disk of third finger which is half again as large as the tympanic area. heels overlap appreciably.—COCHRAN FROGS OF COLOMBIA AND GOIN 307 placed that the light areas between them form a network of fine lines in a reticulate pattern very reminiscent of the pattern of a giraffe. knee and elbow overlap considerably. from Ceilan. its posterior border free with a distinct median notch. truncate in profile. heel reaches almost to tip of snout.5 mm. mm. 18.. 11 mm. head length. roof of skull not exostosed. in postaxillary region Body not dermal appendage on narrower than greatest width of head. rounded patches. Colombia. separated from each other by an interval equal to about from eye. no ulnar ridge..8 mm. a skinfold across chest. H. loreal their distance region flat flaring out hardly at all below it. Skin of head not co-ossified with skull. Canthus rostralis and nearly vertical. no slightly more than one-half webbed. Tympanum distinct. . disk outer metatarsal tubercle. heel. the upper lip slightly defined. by the fact that the latter has at most one or two rows of rounded spots down the middle of the back. Snout moderate. the base of the penultimate phalanx. 10. H. lying close together between the small. femur. no vocal sac. —The entire dorsal surface—including top of head. interorbital distance slightly greater than width of upper eyelid. — head width. Fingers webbed at base. a well-developed narrow glandular ridge encircling upper part of tympanum. adult female. Dimensions. no elongate. web on third finger reaching base of penultimate phalanx. 10. Vomerine teeth in two small. 26 mm. Toes the web on fourth toe reaching third and fifth subequal. palpebral membrane not reticulate. not prominent. reticulata may be separated from its nearest relative. which is about equal the distance between nostrils. 16 mm. laynei.. when hind legs are bent at right angles to body. nostrils more lateral than superior. not particularly projecting. A welldeveloped patagium extends from the back of upper arm to side of body. — MLS 25. Head and body. its diameter slightly greater than its distance from nostril. heel-to-toe. tibia. of fourth just covering the rounded outer but an inner tarsal ridge.. fourth longer than second.. when hind leg is adpressed. 31.8 mm. when limbs are laid along the side. broadly cordiform.

In two of the specimens the dark spots on the back are placed close together so that the light background between them forms a network of narrow lines. laynei can be distinguished at a glance from all other frogs in this group except H. Colombia). In the three specimens in which the facial pattern has not faded the reticulate pattern on the side of the face is quite marked. loreal regions. There is a trace of the pattern on the dorsal margin of thigh but the remainder of thigh is immaculate. and more widely scattered. reticulata by the dorsal dark spots. The pattern in the specimens with the fewest dorsal spots approaches that in H. all from the show much variation in dorsal pattern. same general six vicinity west of Florencia. p. dark spots. and feet is a by large. 61 (type locality. more rounded. CNHM 56198. Florencia. Pucallpa. Amazonas. PERU: Pebas. the dorsal pattern is extremely Variation. It differs . In the other four individuals the dorsal spots are smaller. Goin and Layne. laynei from Leticia. 1958. EPN 2850. 25a. Ucayali Province. Hyla laynei Goin. Hyla laynei Goin Figure 35 1957. Morelia. —The Colombian specimens at hand. Leticia. chocolate brown spots that so pale yellowish gray overlain approach one another that they leave but a reticulated network ground color. reticulata restricted to the lowlands of the Amazon basin and that typical reticulata will be considered a form more characteristic of the eastern slopes of the Andes. SUNHM 6322-3. Lagarto Cocha. Napo-Pastaza: Rio Sandalias. Diagnosis.— NATIONAL MUSEUM BULLETIN 308 U. Remarks. —A small. but we have not seen the type of favosa and indeed do not know if it is extant. EPN 2972. — Specimens Examined COLOMBIA Caqueta: Ceilan.S. although one specimen has a dorsal pattern of rounded. thus. H. 25b. giving the impression of a pale frog with dark spots on the dorsum. discrete. 12621. hands. Loreto Department: Yarenacocha. lower legs. west of Florencia. 2 88 — top of arms. Hyla favosa Cope seems to be a synonym of this species. upper lips. p. All ventral closely of the pale surfaces are a pale cadaverish gray. Based on its description. and that species eventually may prove to be a race of H. west of 25415-7. CNHM 45434. 107. Rfo Ucayali. The dorsal pattern is very similar to that of the reticulated giraffe. reminiscent of the pattern of a reticulated giraffe. brightly colored frog of the leucophyllata group with a pattern of dark rounded spots on a pale brown or tan background. ECUADOR: ANSP MLS 25.

disks of second and third fingers about — equal to size of the tympanum. and 4 webbed to bases of their penultimate phalanges. eyes moderate in size and not pronouncedly protuberant. knee and elbow in contact when limbs are pressed fingers 2.. with a narrow margin of web extending to base of disk of fourth finger. snout somewhat triangular as seen from above with the tip broadly rounded upper jaw projecting but slightly beyond tip of lower. from Leticia. CoDescription. Upper eyelids. UF 8503 (holotype). very slightly notched and slightly free behind its diameter about one-half the width of mouth.—Hyla 337-262—70 21 laynei. Tympanum distinct but small. Figure Zb.. 3. holotype. its width slightly greater than distance from snout to occiput. fingers 3-4-2-1 in order of decreasing length. lombia. . Amazonas. toes 4-3-5-2-1 in order of decreasing length. the distance between the patches about equal to the distance between a patch and a choana. Fingers not particularly slender. Toes slender. UF 8503 (X 2). Vomerine teeth in two small oval patches between the choanae each patch about the size of a choana. diameter of eye about equal to distance from eye to nostril. and dorsum smooth. top of head. nostril near tip of snout. distance from eye to tympanum slightly less than diameter of the latter. Tongue nearly round. interorbital space about equal to diameter of individual eye. not more than one-third the diameter of eye. Head broad. Heels slightly overlapping when legs are flexed with femora held at right angles to the body. FROGS OF COLOMBIA — COCHRAN AND GOIN 309 from that species in having only one or two rows of spots down the middle of the back. fully webbed except for fourth toe which is webbed to the base of penultimate phalanx. Well-developed disks on all fingers and toes.

diameter of eye. 7. In none of the three is the external vocal pouch prominent. — heel-to-toe.7 mm. a single median row of — brown spots extends from between the eyes to just the sacral hump. and top of shank a light tannish gray. Remarks. the shanks.9 mm. Each area is about four millimeters in diameter and lies just posterior to the insertion of arm. 1957. diameter of tympanum. but slightly smaller than. Structurally there is but little variation amongst the three specimens. and the venter was tinged with the same color. Since no additional specimens have been collected since Variation. and thighs were a bright salmon pink. A well-developed patagium. the other darker. In the darkest specimen the ground color is a light brown and consequently the dark brown spots do not show up quite so prominently. NATIONAL MUSEUM BULLETIN 2 88 along the side. 13. Both of the paratypes are 24. 9. 3 mm. These spots are roundish and each is dark.: 310 U. The paired glandular areas do not quite meet on the median line. chocolate posterior to about a millimeter. 10. and continues along the side to groin. in head and body length. heel extending to anterior margin of eye when leg is pressed along body. otherwise the entire ventral surface is immaculate. The hands. Head and body. A dark stripe of the same nature extends from behind each eye through the tympanum and then breaks up into a row of spots similar to. 15. in diameter..5 mm. those on the back. not counting those on top of the head. —The type was collected while the frog was calling in a wet meadow.. head width. 1. The other two do not have the patagium quite so well developed as the type but even in them it is prominent. There is a pair of glandular areas in the skin of pectoral region. — shank. .8 mm. Brown flecks are distributed along the margin of the lower jaw.S. but undersides of thighs and chin nearly smooth. In life this specimen had a dorsal ground color of tan and the under parts of the abdomen and hind legs were salmon pression of a double pink. thus giving the im- row of spots down the back. the two paratypes were taken in grass growing in the . arm. In the specimen with the lightest ground color the dark brown spots are double at two places along the back and between the eyes. Dark spots similar to those in the row along the side occur on top of arms. p. no wellmarked external vocal pouch.1 mm. and the spots were a rich brown.. Venter strongly rugose. Dimensions. series was taken. brownish spots are distributed on the loreal region and upper lip. In life the ground color of the dorsum was yellow Color in lije.. Ground color on top of head. tip of snout to posterior margin of tympanum. dorsum. and rounded. 61) In dorsal ground color one of the paratypes is lighter than the type..6 mm.2 mm. 25. thigh. feet. top of Color in alcohol. and the feet. The upper lip is whitish. or slightly more.8 mm.. we quote the comments on variation from the type — the original description (Goin. In this specimen there are five spots in the middorsal row. and the spots are somewhat smaller than in the other two individuals.

distinct dorsolateral glandular with the fingers virtually unwebbed. the fingers virtually unwebbed. see the discussion of variation reticulata. Hyla hobbsi. in Amazonas. Specimens Examined COLOMBIA Amazon as: Leticia. vomerine teeth usually in a continuous. holotype. related to the single member. — MCZ 28053-7. and MCZ 28074 from Gino-Goje. UF 8503 (holotype). Hyla hobbsi. 22. pencil-like black spots on the ^a Figure 36. on the Rio Apaporis in Vaupes.— FROGS OF COLOMBIA —COCHRAN AND GOIN 311 water at the edge of a cove of a small stream nearby. MCZ 28052 (same size). . The voice a musical. We do not know of any South American tree frogs that are closely folds. hobbsi. —A Hyla with distinct dorsolateral glandular Holotype. rattling trill: For comments on under Hyla is kle-kle-kle-klee. folds. arched series between the choanae. H. hobbsi group Moderate-sized tree frogs with dark. a Paratypes. and the toes nearly completely webbed. collected along the Cano Guacaya. new species Figure 36 — MCZ 28052. There are scattered. Colombia Diagnosis. 8504-5 (paratypes). Colombia. tributary of the Rio Apaporis. with the same data as the type. and with the vomerine teeth in an essentially continuous arched series between the choanae. relationships.

glandular. Body somewhat elongate. Tympanum very distinct. slightly triangular when viewed from above. Skin of upper parts smooth. head width. when limbs are laid along the side. knee and elbow considerably overlap. heel-to-toe. that of belly and lower surface of thigh uniformly and distinctly granular.S. Fingers with just a trace of web at base. nostrils more lateral than superior. Head and body. slightly projecting.. dorsal surfaces. fourth considerably longer than second. separated from eye by a distance equal to about two-thirds its own diameter.. 14. a glandular fold passes above the tympanum and then widens and continues posteriorly to about the level of the adpressed elbows.5 mm. —Vomerine teeth in a single. no tarsal ridge. Snout moderate. Toes almost completely webbed. exostosed. heels meet snugly. NATIONAL MUSEUM BULLETIN and the glandular. its diameter nearly equal to its distance from tip of snout. roof of skull not mate phalanx.. just reaching to disk of third which covers about one-half the tympanic area.7 mm. Skin of head not co-ossified with skull. prominent. heel reaches beyond tip of snout. an indistinct oval inner and a smaller and equally indistinct rounded outer metatarsal tubercle. Eye large. which is relatively wide and slightly greater than distance between nostrils. no vocal sac.. head length. when hind leg is adpressed. third — Dimensions. continuous arranged in a curving row with the convex side anterior and lying between the very large. 17. no projecting rudiment of a pollex. no traces of a skinfold across chest. no ulnar ridge. 21.2 mm. adult female. about three-fifths the diameter of eye. Skin of throat and chest smooth. 312 U.8 mm. nearly rectangular choanae. no dermal appendage on heel. hand. the upper lip flaring out below it. disk of fourth toe covering about one-third the tympanic area. the web on fourth toe reaching middle of penultiloreal region and fifth toes subequal.. femur. 28. broadly rounded. . tibia. No patagium. their distance from end of snout about one-third that from eye. tongue two-thirds as wide as mouth opening. in postaxillary region distinctly narrower than greatest width of head.6 mm. dorsolateral 2 88 folds are nearly black in preservative. the upper jaw extending a little beyond lower. 11. when hind legs are bent at right angles to body.3 mm. We this species is closely related to do not believe that any known black. Can thus rostralis strongly denned concave and oblique. 21. Description oj holotype. 42.. nearly truncate in profile. dorsolateral folds over the shoulders set frog. palpebral membrane not reticulate. its posterior border fused and distinctly but shallowly notched. interorbital distance slightly less than width of upper eyelid.2 mm.. separated from each other by an interval equal to about series two-thirds their distance from eye. The it off at once from all other tree frogs.

numerous. National Museum.5 millimeters (the type) down to 27. widely scattered. dorsolateral light stripe originates on the tip snout and passes posteriorly above the eye and tympanum and terminates just anterior to the groin. In head and body length. as is the type. two on the dorsal surface of each arm. Hylonomus bogotensis Peters (substitute name for Hylonomus.. Colombia). about fifteen scattered over the back.S. 127. 1860 [Amphibia. three individuals (MCZ 28053-4 and 28074) are pale gray dorsally. preoccupied). a teeth behind level of or external evidence of ear. and two on the left foot and one on the right Each pale. of the Variation: —Among the six paratypes. In the pale specimens these stripes are hardly apparent except where they pass through the dark glandular.of the U. Bogota. Jr. Hobbs. pencillike dots on the dorsal surface. Diagnosis. four on the left shank and three on the right shank.8 millimeters in an immature individual. In the darker specimens. type species. and pale dorsolateral stripes. p. Hylonomus bogotensis Peters (preoccupied by Hylonomus Dawson. 107 (type locality. Hylonomus bogotensis Peters. rounded median subgular vocal pouch in male. FROGS OF COLOMBIA Color in alcohol. the specimens range from 42. In all except one of the dark specimens (MCZ 28056) are apparent pale dorsolateral stripes that originate at the tip of the snout and pass above the eye and tympanum and terminate in the region of the groin. however. Stegocephalia]) Hyloscirtus Peters. for the six specimens in the series (MCZ 28052-7) were all collected and preserved by the same collector at the same time and place. The pattern — Ground —COCHRAN AND GOLN 313 above and below. dorsolateral fold. This variation in intensity of dorsal color probably is not due to preservation alone. The other three specimens are much darker dorsally. vomerine choanae. . three on the dorsal surface of each thigh. —We dedicate this species to Dr. Genus Hyloscirtus Peters 1882. There are three dorsal dots on each loreal region. Hyloscirtus bogotensis (Peters) Figure 37 1882. 107 (type species. Hylonomus Peters. this light stripe stands out distinctly against the dark background. narrow. p. three on top of the head. Life history unknown. dark. 1882.. foot. These stripes are very indistinct on this specimen because of its pale ground color. Horton H. p. above the color very pale gray consists of three elements: the glandular folds shoulders. which are black. Remarks. —No tympanum sacral diapophyses. although close inspection shows that the same basic dorsal pattern is present in all of the specimens.

male (at female (both same size). — Noble. 1944c. bogotensis. from the species of Hyla it may be distinguished by the absence of a distinct tympanum. . 1882.S. slightly projecting. and from MLS 160. Vomerine teeth in two transverse series which are slightly more anterior medially. lying close together behind the level of the small. 1917. Snout moderate. its posterior border slightly free and slightly notched. preoccupied) Dunn. — 1963. 1923. p. p. somewhat U-shaped when viewed from above. rounded choanae. tip somewhat rounded in profile. by having the diapophyses of the sacral vertebrae essentially rounded. Description. p. 1882. their distance Hyloscirtus bogotensis: CNHM 81916. 398. p. p. left). and by usually having the vomerine teeth in transverse series behind the choanae.— — NATIONAL MUSEUM BULLETIN 288 314 U. —A hylid tree frog with rounded sacral dispophyses. 516. and the vomerine teeth in transverse series which usually lie well behind the posterior margins of the small. — Peters. Hyloscirtus bogotensis. 1944a. Cundinamarca. 12. Diagnosis. 23. tongue three-fourths as wide as mouth opening. p.— Gorham. p. — MLS 160.— Goin. Figure 37. from Chipaque. an inconspicuous tympanum. Hyloscirtus bogotensis may be separated from the species of Cryptobatrachus by the absence of a distinct tympanum. Colombia. rounded choanae. broadly cordiform. the upper jaw extending beyond lower. nostrils lateral. 1961. 127 (substitute for Hylonomus Nieden. 805. 73. .

. in postaxillary region a little narrower than greatest width of head. no projecting rudiment of a poll ex. skin of throat and chest smooth. Dorsal ground color a pale grayish brown rather thickly sprinkled with pepper-like black specks. Head and body. when limbs are laid along the side..9 mm. Tympanum slightly distinct. hand.. 21 mm. 14..FROGS OF COLOMBIA —COCHRAN AND GOIN 315 end of snout about one-half that from eye. 22. 36 mm. Skin of head not co-ossified with skull. Color in life. disk of fourth toe slightly greater than the tympanic area. no ulnar ridge.3 mm. head width.. Toes slightly more than one-half webbed. Fingers webbed at base between fingers 3 and 4 and with a trace of webbing between fingers 2 and 3. interorbital greater than width of upper eyelid. that of belly and lower surface of thigh uniformly granular. femur.7 mm. when hind legs are bent at right angles to body. Color in alcohol.. head length. separated from each other interval equal to about their distance from eye. just reaching to disk of third which is about one-third again as great as the tympanic area. knee and elbow fail to meet.4 mm. about one-third the diameter of eye. its diameter equal to its distance from nostril. 14. which distance slightly about equal the distance between nostrils. by an . The specks are reduced on the top of head and snout and are less prominent on the dorsal surfaces of thighs and forearms than they are on the dorsal surfaces of the rest of the limbs. the web on fourth toe reaching base of penultimate phalanx. 48. Skin of upper parts smooth. There was some evidence of metachrosis — — is . All ventral surfaces were a dirty gray without pattern except for faint pepper-like flecks under the throat and a yellowish tinge to the under-margin of the jaws. Dimensions. in life. no tarsal ridge. tibia. then curving downward to above insertion of arm. when hind leg is adpressed. the upper lip flaring out but slightly below it. 15. a heavy glandular fold passing above upper part of tympanum. Eye neither large nor prominent. a yellowish brown with a definite pattern of more or less uniformly distributed pencil-like black spots on all the dorsal surfaces. separated from eye by a distance nearly equal to its own diameter. A trace of a patagium extends from the back of upper arm to side of body. adult female. a distinct oval inner but no distinct outer metatarsal tubercle.7 mm. heel reaches posterior margin of eye. collected by Alice Hunter) was. fourth finger about one-half disk longer than second. no vocal sac. heels overlap greatly. palpebral membrane not reticulate. Body rather elongate. very faint traces of a skinfold across chest. third and fifth toes subequal. An adult female (CJG 2299. no dermal appendage on heel. roof of skull not exostosed. Canthus rostralis rather well defined loreal region slightly concave and nearly vertical. heel-to-toe. where it terminates. The entire ventral surface is a cadaverish gray with faint traces of flecking on the chin and throat.

the color pattern variation seems to be a result of sexual dimorphism. just as the adult females. vicinity . the one male specimen (CNHM for the type. MZUM 123946-8. Dr. The immature female (CNHM 54776.316 U. 46 mm. for it is Specimens Examined COLOMBIA Cauca: El Tambo. and Mrs. an adult female (MLS 160) from Chipaque. Bogota. (CNHM 81916) from Boquer6n Gorge. Chipaque. CNHM 54776.) generally darker and has the dorsal pigment concentrated into rounded black spots that are scattered over the upper surface Dunn described for the male he collected. All of the adult females we have seen are grayish yellow with tiny. 81916. nine specimens of this species have been collected: the type. Cundinamarca: Boqueron Gorge. just as Dr. for the glass jar than — scattered black spots above. an immature female (CNHM 54776) from El Tambo in Cauca. We have seen seven of the nine known specimens. The conspicuousness function of of size. an adult male. Remarks. Niceforo Maria in the vicinity of Bogota. 1962. To our knowledge. Dunn for his male. Furthermore. One adult female (MLS 160) contains what are apparently mature eggs ready for desposition. As Dunn suspected. just as On is was described the other hand. Further studies on the life history of this species may throw further light upon its relationships. an adult female (CJG 2299) collected by Alice Hunter in Boquer6n Gorge. The only specimen with known ecological data is the one collected by Dr. Dunn. Bogota. Bogota. and two adult females and a juvenile (MZUM 123946-8) collected by Hno. NATIONAL MUSEUM BULLETIN 2 88 specimen became paler after sitting for a few minutes in a it had been when taken from the collecting bag. 1944. Dunn near Bogota on July 21. 33.) is pigmented larger. Cundinamarca. a male collected by Dr. The structure of the girdle may well imply a relationship with Cryptobatrachus. described by Peters in 1882. although it certainly is not as conspicuous as it is in Cryptobatrachus. If the eggs are unpigmented when desposited it would suggest they are deposited in a secluded site. the tympanum is apparently partly a not in evidence in the immature female but is discernible in both adult specimens.S. 160. this male specimen has a round dark spot under the chin as described by Dr. CJG 2299. CNHM MLS 81916. of Bogota. but no pigment is discernible on them. January 8. Apparently these differences are purely sexual.6 mm. but whether or not the eggs are carried on the back is unknown. Dunn suggested that it might breed in bromeliads and that its tadpoles are modified for that habitat. it was taken from a bromeliad that had grown to about four feet.

broadly cordiform. 69710. p. Ecuador. Hyla orcesi Funkhouser. Huila. a2 O. roof of skull very weakly to markedly exostosed. 38. and lack of large exostoses on the skull. Canthus rostralis rather well defined. may be distinguished from 0. Description. spiny warts giving the back a sandpapery top of skull heavily exostosed O. taurinus by its smaller size. its posterior border slightly free and shallowly notched. broader than long.)' 1963. more or less transverse series.FROGS OF COLOMBIA —COCHRAN AND GOIN 317 Genus Osteocephalus Steindachner 1862. . effect. the upper lip flaring out only slightly below it. tongue three-fourths as wide as mouth opening. nostrils their more lateral than superior. Generic diagnosis. orcesi like —CNHM lower. Osteocephalus orcesi (Funkhouser) Figure 1956. Plate 38a-c 78 (type locality. near San Adolfo. Osteocephalus Steindachner. top of skull moderately exostosed O. 22. 77 (type species. Vomerine teeth in two short. Osteocephalus taurinus Steindachner). skin on top head neither thickened nor co-ossified with skull. rounded head. Dorsum with numerous. —A small Osteocephalus with numerous spiny warts on the back which give the dorsum a sandpaper-like effect. orcesi b l . the upper jow extending very little beyond 0. nearly truncate in profile. ovate choanae. Ieprieuri Top of head rounded and not heavily exostosed. considerably projecting. p. rounded when viewed from above. to Colombian Species of Osteocephalus head flattened and moderately to heavily exostosed. loreal region concave but not particularly oblique. interorbital distance slightly less than width of upper eyelid. Suno drainage. . from Acevedo. Diagnosis. Snout moderate. 600 m. small. Ieprieuri by the sandpaperback and from 0. not prominent. palpebral membrane not reticulate. Rio Suazo. canthus of rostralis nearly straight. Colombia. tributary of Rio Cotapino. p. Eye moderate. lying close together between the moderate. canthus rostralis curved. Province. is relatively wide and slightly greater than distance between which nostrils. taurinus Dorsum smooth or slightly roughened but not with pronounced sandpapery b2 effect. distance from end of snout about one-third that from eye. — Males with paired vocal pouches. Napo-Pastaza — Gorham. separated from each other by an interval equal to about two-thirds their distance from eye. of Key a Top 1 . . its diameter equal to its distance from nostril.

Toes more than one-half webbed. disk of fourth toe covering about two-thirds the tympanic area. lap heels overlap considerably. Skin of upper parts of head. heel reaches beyond tip when limbs are laid along the side. with a few less-distinct warts on the loreal region. skin or throat and chest smooth. of snout. that of belly and lower surface of thigh uniformly granular.S. NATIONAL MUSEUM BULLETIN 2 88 Tympanum very distinct. . A rather narrow glandular ridge encircling upper part of tympanum. reaching to middle of disk of third which covers about three-fourths the tympanic area. conical. when hind leg is adpressed. no tarsal ridge. fourth finger considerably longer than second.— 318 U. CNHM69 710 (same size). adult male. in postaxillary region a little narrower than greatest width of head. web on third finger reaching base of penultimate phalanx. when hind legs are bent at right angles to body. the web on fourth toe reaching base of penultimate phalanx. No patagium. Osteocephalus orcesi. no projecting rudiment of a pollex. separated from eye by a distance equal to three-fifths its own diameter. white-tipped warts. Fingers webbed at base. third and fifth toes subequal. legs. knee and elbow overconsiderably. Figure 38. and feet thickly covered with small. no traces of a skinfold across chest. about three-fifths the diameter of eye. no ulnar ridge. Body mod- erately elongate. See also Plate 38a-c. trunk. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. no dermal appendage on heel.

Brazil). 94. 1927. Rio Pastaza watershed). p. . 1923. p.— De Witte. hand.— Boulenger. 22. 1961. spots along each side between axilla and groin. 13150 (type). roof of skull weakly exostosed. Oriente. p. Osteocephalus flavolineatus Steindaehner. p. 637. A rather distinct series of black. 1904. Rfo Negro. p. 27619. but on direct comparison with the type of britti the two proved to be different. and legs. Peracca. head length.. 1862. 64. 1931.6 millimeters head-body length. 64. Hyla taurina.- — — — A large Osteocephalus with the skull strongly exostosed and with numerous small. 9. p. p. 1941. p.—When we first examined one of the specimens we felt sure we had Melin's Hyla britti at hand. 1867. p. 13.— Lutz. ECUADOR: SUNHM CNHM 25791.1 mm. 1951. Brazil). 48 mm.. Rio Pastaza basin. 1898. Hyla taurinus. p. tributary of Rfo Cotapino.— Crawford.— Ruthven. 1935. 447 Lutz and Kloss. 133. wart-like spines on the back which give the dorsum a sandpaper-like effect. 27 mm. p. Abitagua. Rio Suaza.— Melin.3 to 54. 1952. Cocuy. p. dirty brown. Miranda-Ribeiro. 35. 1912. 1951.— Beebe. p. p. probably valid. Trachycephalus flavolineatus. Diagnosis.3 and body. 38. Suno drainage. 38. 1919. 14. 1867. Remarks. heel-to-toe. — — 1904. Ecuador. near San Adolfo. —Head 17. 1930. 14. p.— Rivero. Napo-Pastaza Province. 208. mm. — — Trachycephalus taurinus.— Baumann. Entire ventral The numerous white-tipped warts on the dorsum and top of legs give this individual a speckled appearance. 37.— Nieden. 73 (type locality. 1945. Skin of head not co-ossified with skull. 1919. 303. femur. 1898b. 1882a... Gorham. p. After studying the type of Hyla orcesi (SUNHM 13150) we have no doubt that the specimens at hand are orcesi and that britti is a different. p. Steindaehner. 163. p.7 22. Osteocephalus taurinus Steindaehner Plate 38d-f 1862. 33. p. species.. 654. p. p. p. 1963. The in five specimens we have seen range from 47. 1884a. p. head width. —Very dark brown above with indistinct scattered Color in alcohols blotches of deep chocolate brown. 80 (type locality. 77 (type locality.7 mm. 17 mm. 1926. Indistinct crossbars on upper surfaces of arms surface a dull. Steindaehner.— Melin. Dimensions. Specimens Examined COLOMBIA Huila: CNHM Acevedo. 363. Hyla depressa Andersson.— Parker. 110. 1867.— Schmidt and Inger. 512.FROGS OF COLOMBIA COCHRAN AND GOIN 319 vocal sac at each angle of jaw but apparent only on dissection. 1961. 69709-10.— Boulenger. tibia. p.— Goin. sometimes ocelli-like. 1867. 40. mm.. p. Osteocephalus taurinus Steindaehner.

Serrania de La Macarena. and from 0.distance from eye. slightly projecting. no dermal appendage on heel. Fingers webbed at base. Toes more than three-fourths webbed. its posterior border fused and unnotched. the upper jaw extending slightly beyond lower. web on third finger reaching middle of antepenultimate phalanx. small spines on the back. their distance from end of snout about one-third that from eye. heels overlap greatly. NATIONAL MUSEUM BULLETIN 288 may be distinguished from 0. nostrils more lateral than superior. heavy glandular ridge passing above tympanum. a few less-welldeveloped tubercles on upper surfaces of arm and shank. rounded. Skin of upper parts glandular. with numerous. loreal region concave but nearly vertical. not particularly prominent. somewhat pear-shaped. third and fifth toes subequal. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Two very prominent. separated from each other by an interval equal to about two-thirds then. roof of skull heavily exostosed. one behind each angle of jaw. adult male. the upper lip flaring out but slightly below it. Eye large. . lying between the large oblong choanae. its diameter equal to its distance from nostril. no tarsal ridge. tongue one-half as wide as mouth opening. Tympanum very distinct. no ulnar ridge. disk of fourth toe nearly covering the tympanic area. Vomerine teeth in two heavy /~~\ series. interorbital distance slightly greater than width of upper eyelid. a pronounced. knee and elbow overlap considerably. small. truncate in profile. skin of throat and chest smooth. Snout moderate. when limbs are laid along the side. longitudinal ridges on top of head between the eyes. fourth finger considerably longer than second. Body rather elongate. with a pair of well-developed longitudinal ridges. vocal sacs paired. palpebral membrane not reticulate. no projecting rudiment of a pollex.S. just reaching to middle of disk of third which just covers the tympanic area. heel reaches well beyond tip of snout. in postaxillary region a little narrower than greatest width of head. Skin of head not co-ossified with skull. Can thus rostralis sharply defined. which is relatively wide and slightly greater than distance between nostrils. that of belly and lower surface of thigh uniformly granular. wart-like tubercles. when hind leg is adpressed. half again as long as wide. no traces of a skinfold across chest.320 U. Description. from Meta. taurinus of heavily exostosed skull. No patagium. about three-fifths the diameter of eye. leprieuri by the presence numerous. orcesi by the 0. U-shaped when viewed from above. —CNHM 81332. the web on fourth toe reaching the middle of penultimate phalanx. when hind legs are bent at right angles to body. separated from eye by a distance nearly equal to one-fourth its own diameter.

. USNM BM 118057. The specimen from Chenapoka is heavily peppered with brown on throat and chest. Ecuador.119.8 Color in alcohol. Brazil. 64) called jlavolineatus and Andersson (1945. There are faint vermiculations of grayish brown under the chin. Colombia.. 43 mm.. CNHM 69711-4. p. 1905. Kartabo. 1878. 21. This is what Steindachner (1882. BOLIVIA: Buenavista. The specimen (BM 1927.12. MCZ 2S060.3 mm. ZMU 487 (2).16. 52 mm. is from Chenapoka. the largest specimen of taurinus we have seen (BM 1905. femur. Meta: Serrania de La Macarena. —A very dark brown frog with just a hint of darker blotches on the back.8. Gino-Goje. Remarks. 23. type of depressa. 73) called depressa. Dr. We have no evidence that any of these variants represent geographic races worthy of nomenclatorial recognition. 7 specimens. heel-to-toe. Buenavista. shanks. BM ZMU 487 seen..11.1. tibia. these frogs have green bones. Demerara. BM 1927.1. CNHM 81332. aflucnte del Cubigu. BM . Three fairly distinct broad crossbands can be seen on each thigh with less-distinct broad crossbands on arms. mm.20-21. in life. p. Fred Medem has pointed out to us that. FM Vaup6s: Caiio Negro. Putumayo: Rio Mecaya. British Guiana.1 mm. know these frogs by the common name "mamoen" and use them as food. BRAZIL: Rio Vaupes. At the other geographic extreme.— The Indians in Vaupes. and (2). hand.119.11. Variation. 37. Rio 1927.4 mm.4 mm. Bolivia.18.2.FROGS OF COLOMBIA —COCHRAN AND GOIN 321 — Dimensions.13. The vocal sacs are black.20). and — taurinus.. has a gray rather than brown ground color and in this respect differs from all other specimens of taurinus we have seen four specimens Pastaza. BRITISH GUIANA: Chenapoka. Specimens Examined COLOMBIA Amazon as: Rio Apaporis.1.Rio Vaupes. measuring 93 millimeters in headbody length..1.8. Bolivia) from the western part of the range of taurinus that have a conspicuous yellow stripe down the back. 8.119) from Buenavista. There is quite extensive variation in specimens of 0. head length. BM 1956. (unnumbered). 72. while most western specimens are immaculate or very weakly mottled on those areas. MCZ 28048. Head and body. We have (NR collection. The ventral surface is a dirty brownish gray. Lower Rio Apaporis. We have seen no specimens from the western part of the range that reach this size. head width.1.1. 22. feet.

1941. Osteocephalus leprieuri (Dumeril and Bibron) Plate 39a-c 1841." EPN 1679-80. Osteocephalus planiceps Cope. Cope. Tympanum very it. Rio Rutuno.12. Brazil. 1919. the upper lip flaring out moderately below Eye neither large nor prominent. 81. Hypsiboas 1874. which is relatively wide and onethird again as great as the distance between nostrils. p. MCZ 28042. leprieuri.63. p. p. 1961.— Beebe. USNM 83947. Rio Pastaza. CNHM 39853. p. EPN 4039. 23. Taracua. NR collection (type of Hyla depressa). p. French Guiana.S.— 322 U. p. 22. 1882. Moruna-Santiago. p. Hyla vilarsi Melin. its diameter equal to its distance from nostril. PERU: Ayacucho. leprieuri may be distinguished from 0. tosed skull and a relatively smooth dorsum. EPN 133G-9. 40 (type locality. 200. NATIONAL MUSEUM BULLETIN 2 88 ECUADOR: Macuma. p. EPN 1347-8. Osteocephalus leprieuri. VENEZUELA: Cerro Ya-Pacana. — 1961. Hyla Nieden. loreal region concave and oblique. 122 (type locality. 101. EPN 1522. p. Hyla planiceps—boulenger. from Vaupes. 13. 1904. EPN 1344. p. somewhat rounded when viewed from above. 1961. p. EPN 1430. Snout neither large nor prominent. nostrils decidedly lateral than superior. Cayanne). Nauta). Peru. taurinus by having the dorsum smooth instead of sand-papery.— Rivero. rather pointed in profile. Huancahamba. leprieuri Dumeril and Bibron. Rio Corrientes. Pindo. 306 Gorham. 1874b. in two moderately heavy. oblong choanae.— Nieden. tongue three-fourths as wide as mouth opening. p. 1923. — the upper jaw extending considerably beyond lower. Montalvo. p. 1882a. 208 — Miranda-Ribeiro. EPN 2502-4. 1923. 3811-28. p. cabeceras del Rio Capahuari. Rio Vaup6s).— Baumann. 361 1912. p. Colombia. 0. lying very close together between the rather large. 553 (type locality. Vomerine teeth Description.13. p. Lidth. 1867. transverse series. Rio Pucayacu. 305. EPN 1393. EPN 2582. — — — Andersson. Sivia on Rio Apurimac. —A medium-sized Osteocephalus with a moderately exosDiagnosis. 94. separated from each more other by an interval equal to about two-thirds their distance from eye. p. p.— Boulenger. interorbital distance about equal to width of upper eyelid. 1958. . 364. 1882a. broadly cordiform. Gorham 1963. orcesi and 0. 73. BM 1911. Goin. 163. — 1961. considerably projecting.— Gunther. 1926. mas abajo de Montalvo. Canthus rostralis very strongly defined and practically straight. 1945. p. palpebral membrane not reticulate. "Region de Puyo. its posterior border slightly free and very shallowly notched. 13. Rio Pindo. 130. their distance from end of snout about two-fifths that from eye. 1867. — Goin. 1963. Napo-Pastaza.

through eye and tjmipanum to fade out above the insertion of arm.. we would . head width.2 mm. rather distinct dark bands. Body rather elongate. No patagium.1 mm.. 38. Head and body. heel-to-toe. nominal forms synonymous with it is it. planiceps. 54. 18. — Dimensions. 17. Skin of upper parts rather smooth. 31. no vocal sac. about four-fifths the distance equal to one-half its appendage on heel. The specimen described and illustrated (MCZ 28042) has been directly compared with the types of leprieuri. no dermal distinct. and behind this are a couple of even less distinct transverse bars across the back. 17. third and fifth toes subequal. heel reaches tip of snout. the web on fourth toe reaching base of penultimate phalanx. —Dorsum dark brown. and.9 mm. and vilarsi by the junior author and there seems to be no doubt that all are conspecific. reaching to middle of disk of third which covers about one-half the tympanic area. in postaxillary region a little narrower than greatest width of head. tibia. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. — Remarks. The arms.. hand. A very dark brown (nearly black) facial stripe runs from the tip of snout across the loreal region. knee and elbow overlap considerably. no tarsal ridge. femur. a distinct but narrow glandular ridge passing above tympanum. are considered conspecific. disk of fourth toe covering about onethird the tympanic area. Adult gravid female. and The entire ventral surface feet are each crossed is with broad. when limbs are laid along the side.FROGS OF COLOMBIA — COCHRAN AND GOIN 323 diameter of eye. when hind leg is adpressed. Toes more than one-half webbed. not unlikely that there are other although we have not as yet seen types of any material other than those mentioned above that assign to the synonymy at the present time. when hind legs are bent at right angles to body. skin of throat and chest smooth. fourth finger considerably longer than second.2 mm. Another specimen (CNHM 697 1G) has been directly compared with the types of planiceps and vilarsi and these. Fingers webbed at base. indeed.5 mm. no ulnar ridge. Skin of head not coossified with skull. no projecting rudiment of a pollex. roof of skull moderately exostosed.. that of belly and lower surface of thigh uniformly and very finely granular..3 mm. traces of a skinfold across chest. another somewhat bilobed one above the shoulders. head length. likewise. heels overlap appreciably. 27. It is not surprising that this wide-ranging species has been described at least three times.. a cadaverish gray with dusky mottlings present on the lateral portions of chin and throat. Dorsally there is an indistinct dark blotch on top of the head. separated from eye by a own diameter. mm. legs.1 Color in alcohol.

cauda de La Playa. EPN 1387-89. PERU: Nauta. 1961. large. 22 (type locality. 152139. 23. Rio Villano. 152203. Venezuela. ECUADOR: Napo-Pastaza: Montalvo. Department of Zulia. USNM MLS 16. Pindo. p. Rio Apaporis. The —A ingens.S. — Rivero. Phrynohyas ingens Duellman. uniformly colored Phrynohyas with a broad Phrynohyas venulosa at hand has a headand a head width of 27. ANSP 11399 (type. male with paired.5 millimeters — . broadly rounded. from Astillero. — p. with or without a dorsal pattern. Phrynohyas ingens Duellman Plate 39d-f 1956. 69176. BRAZIL: Rio VaupSs. USNM CNHM USNM MCZ Vattpes: Gino-Goj6. tongue four-fifths as wide as mouth opening. —The 30 (type species." Putumato: Rio Mecaya. Hyla tibiatrix Hyla venulosa Diagnosis. p. lateral vocal pouches. Hyla zonata Spix). 152010-1. p. vomerine teeth in two short series between the rather small. Taracua. 1963. 11. no postorbital process on frontoparietal bone. head. Vomerine teeth in two short.3 millimeters. 448 (type. NATIONAL MUSEUM BULLETIN 2 88 Specimens Examined COLOMBIA USNM Amazonas: Leticia. and with a rather narrow head (head width generally less than 30 mm. 1961.5 and 94. Moderate-sized. 152136-8. Norte de Santander. 14. uniformly colored.5 millimeters. Colombia. EPN 1519-20. no precise locality. Goin. EPN 1341-42. Phrynohyas Fitzinger. 1961. La Fria). nape. Generic diagnosis. leprieuri). vilarsi). 28042. Cabeceras de Rio Capa huari. rounded choanae. 1963. MCZ 28050. rounded choanae. EPN 1488-9. Key a a2 to Colombian Species of Phrynohyas Large. Meta: "El mico. p. Rio Apaporis. Puerto Narino. ZMU MHNP FRENCH Genus Phrynohyas Fitzinger 1843. GUIANA: Cayenne. heavy. Gorharn. 131. — 1963. planiceps).6 millimeters and head widths of 31.) P.324 U.) 1 . lying between the moderate-sized. MLS 265a. frogs of this genus show the following combination of characters: skin of head not co-ossified with skull and roof of skull not exostosed. the two specimens of P. ingens. Description.8 and 33. — Rivero. ingens greater than 30 mm. its posterior border free largest specimen of body length of 84. and with a broad head (head width generally P. 4629 (type. transverse series. p. Pueblo Nuevo. skin on top of head. venulosa . and shoulder region thickened. ingens at hand have head-body lengths of 94.

skin on top of not co-ossified with skull. when hind legs are bent at right adpressed. Skin of upper head and shoulders thicker than skin on back. fourth finger a antepenultimate third little longer than second. and lower surface of thigh uniformly granular. not prominent..5 mm. no projecting rudiment of a pollex. 42. belly. head length. 94. skin of throat. Toes slightly more than one-half webbed. but slightly projecting. axillary region about equal greatest Body rather elongate. heels overlap slightly. traces of a skinfold across chest. rounded in profile. 31.8 mm. knee and elbow side.8 mm. interorbital distance about equal to width of upper eyelid. no dermal appendage on heel. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. Eye neither large nor prominent. when viewed from above. no tarsal one-half its of ridge. ingens. ventral surfaces a cadaverish gray-brown. roof of skull not exostosed. Fingers webbed at base. Dorsum pale brown.7 mm.. 29. a distinct glandular ridge passes down at an angle over the tympanum and towards the insertion of arm. palpebral membrane not reticulate. web reaching distal phalanx of third finger. their distance from end of snout about two-thirds that from eye. heel-to-toe. separated from eye by a distance nearly equal to and with just a hint of a notch. third and fifth toes subequal. no ulnar ridge.-— Head and body. 46. P.FROGS OF COLOMBIA —COCHRAN Snout AND GOIN 325 rounded upper jaw extending but little beyond lower. nostrils more lateral than superior. femur. which is slightly greater than distance between nostrils. 24 mm. heel reaches almost to eye. At first glance this frog seems to be absolutely uniform in color above. chest. separated from each other by an interval equal to about three-fourths their distance from eye. Skin of head angles to body. reaching to middle of disk of third which just snugly covers the tympanic area. This large species seems to be endemic to the Maracaibo Basin. Dimensions. fail to No patagium. parts somewhat glandular. Remarks. disk of fourth toe covering about two-thirds the tympanic area. when hind leg is when limbs are laid along the meet.. Color in alcohol. however. does — — 337-262—70 22 . adult female. The two specimens recorded here are the first to be taken in the portion of the basin belonging to Colombia. Canthus rostralis not strongly defined.. the web on fourth toe reaching the middle of penultimate phalanx.. the own diameter. its diameter equal to its distance from nostril. the upper lip flaring out distinctly below it.. but on close inspection rather widely spaced wart-like pustules on the back can be seen to be darker in color than the rest of the dorsum. tip slightly short. no vocal sac.2 mm. Tympanum very distinct. head width. in post- width of head. hand. 64.2 mm. loreal region concave and somewhat oblique. tibia. about three-fourths the diameter of eye.

1956. 12. p. 1926. —Peters.5 millimeters. 1961. — Rivero. Peters. 33.— Cott.— Nieden. 58. 1841. 1933. 1952. 326 U. — Dumeril and Bibron. 112. p.— Boettger. p. p. 120.— Gorham.5 millimeters. 1901. p. 35. 37. Spix. p. Astillero. 85. Phrynohyas zonata.— Duellman. p. p. Hyla venulosa venulosa. 31 (type locality restricted by Duellman to TefT6. p.— Santos. p. the smaller of hand has a head-body length of 94. p. 711 (type locality. 53. p. 563 (type locality. p. 103. Hyla tibiatrix tibiatrix. 174. 1855. . 367. 244. 133. 1856. 89. p. North 1858. p.S.— Peracca. 1881.— Gorham. — Lichtenstein. 1923. 1958. —Baumann. 1952. — Guichenot. 1882. 1912. Suriname?). of moderate its size. —A Phrynohyas with or without a dark central patch on P. Brazil). Hyla 1867.— Peters. venulosa may be distinguished from P. p. p. p. 219. 40. venulosa. Brazil). 135 (type locality. p. the dorsum. 1964a. 1912. 102. 1955. 1898b. p. p. p. 23. 208. 1872. Amazonas. 102 (type locality. as three of the specimens of venulosa that we have seen from Colombia are from Astillero. 1S66. 1919. —Boulenger. 1961. — Cope. at the mouth of Rio Teffe' where it empties into the Rio SolimSes.— Proctor. 1952. Lutz. 1961. p. 799. p.— Nieden.— Roux. Hyla 84. 1942. p. 14. p. 302. 6. p. Hyla lichenosa Gunther. 655. p. 76.— Miranda-Ribeiro. America?) p.— Stejneger. p. 1923. p. — Rivero. — Gtinther.— Lutz and Kloss. 364. 1907. 127. 302. p. 64— Niceforo Maria. 1841. 1960. 180. Specimens Examined COLOMBIA Norte de Santander: MLS 265. 1892. 1867. 1873. 265a. Hyla bufonia Ecga [=Ega].— Baumann. ingens— the only other species Phrynohyas presently known to occur in Colombia—by Diagnosis. p. 1S58. 1921. 1952. 1964b. p. 105.— . p. Amazonas. 560. Hyla resinifictrix Goeldi. —Brocchi. p. 1904. Lago 1824. 1923. p. —Taylor. p. 1866a. South America). Hyla vermiculata Dumeril and Bibron. p. Acrodytes venulosa. venulosa. p. p. Rana venulosa Laurenti.— Boulenger. p.— Gorham. Goin. p. p. 42 (type locality. 1894. — Steindachner.— Beebe. 1926. Scytopis venulosus. p.— Parker. 1963.— Gans. 189. 219.— Nieden. 97. 1882a.— Cochran. p. 1927.— Mole and Urich. p. 1164. p. NATIONAL MUSEUM BULLETIN 2 88 not occur in the Maracaibo Basin to the exclusion of P. 303. 1930. p. hand having a head-body length the two specimens of ingens at the largest specimen at of 84. 1964. 1872a. 771. 41 (type locality. 311. 1841. Brazil). coriacea Peters. p. p. p.— A. Basin). 309. 1963. p. 1958. — p. Para. Hyla zonata Amazon Spix. Phrynohyas venulosa (Laurenti) Plate 39g-i 1768.— Goin and Layne. 62. 1824. 1963. 21. 104. 1926. 1887. the same locality that our two specimens of ingens are from. 37. 1956. 23.

28. its diameter equal to its distance from nostril. the upper jaw extending slightly beyond lower. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. 72. it then begins to taper and Color in alcohol. when hind legs are bent at right angles to body. somewhat projecting. its posterior border slightly free and very shallowly notched. equal to their distance from eye.5 mm. in postaxillary region about equal greatest width of head. 22. when limbs are laid along the side. not prominent. Skin of upper parts glandular. Fingers webbed at base. somewhat rounded when viewed from above. femur. The dark pattern consists of a broad band that originates between the eyes. heel reaches posterior margin of eye. about threefourths the diameter of eye. separated from eye by a distance equal to about one-half its own diameter. the upper lip flaring out a little below it. web reaching middle of antepenultimate phalanx of third finger. 34.. no ulnar ridge. 53.7 mm. broadly rounded in profile. transverse series. no dermal appendage on heel. knee and elbow slightly overlap. heel-to-toe. belly. the web on fourth toe reaching base of penultimate phalanx. concave and slightly oblique. no tarsal ridge. Norte de Santander. interorbital distance about equal to width of upper eyelid. adult female. Colombia. wart-like tubercles scattered over the posterior portion of the back. tibia. roof of skull not exostosed. the thick. traces of a skinfold across chest. no projecting rudiment of a pollex. head length.3 mm. chest.. rounded choanae tongue threefourths as wide as mouth opening. separated from each other by an interval . Description. — Dimensions.8 mm. Head and body. from Astillero.. Eye small.. Tympanum very distinct.5 mm. nearly round. with small. third and fifth toes subequal. Can thus rostralis slightly defined. rounded. skin of throat. Snout moderate. their distance from end of snout about two.. when hind leg is adpressed. hand. . extending down on the sides. Body not elongate. glandular skin on top of the head lapping over the upper part of tympanum. passes posteriorly in a slightly sinusoidal curve to the region of the sacral hump where it abruptly broadens. palpebral membrane not reticulate. heels overlap very slightly No patagium.7 mm. 25. fourth finger about a disk longer than second. no vocal sac. disk of fourth toe covering about four-fifths the tympanic area. Skin of head thickened but not co-ossified with skull. and lower surface of thigh loreal region coarsely granular. reaching to middle of disk of third which just covers the tympanic area. nostrils more lateral than superior. Vomerine teeth in two short. Toes three-fourths webbed.thirds that from eye. which is about equal to the distance between nostrils. —Dorsal ground color light brown with a superimposed pattern of dark brown. lying close together between the moderate-sized.FROGS OF COLOMBIA — COCHRAN AND GOIN 327 — MLS 258a.. head width. 38.4 mm.

— Rivero (1961. the International Zoological .328 U. In addition to these six available names. The ventral ground color is about the same tone of brown as the dorsal ground color. in head-body length) but are also more slender. and feet. A few have faint. Some have indistinct dark crossbands across the legs. when — 56. (For a study of the variation outside of Colombia. we suspect. The remaining ten specimens from Colombia now before us are stoutmore or less uniformly brownish frogs. NATIONAL MUSEUM BULLETIN 288 terminates above the vent. together with distinct bands across the thighs.0. a standard deviation of 2. 1768 (as published in the combination Hyla tibiatrix) and that it Nomenclature. small spots on the dorsum. The number of maxillary teeth on the left side of our four specimens from Leticia ranges from 66 to 78. We note three types of variants in the Colombian material at hand. others show no evidence of such markings. Goin and Layne bodied. with a mean of 54 and of maxillary teeth between specimens from bia. These apparently are what Dumeril and Bibron called Hyla vermiculata. bujonia. The thigh.65. shanks. transverse. there are several other forms now masquerading as Hyla which. solution today than it was when Duellman (1956) published his study. (1958. and 42. we refer the reader to Duellman's report. Two specimens from Astillero and one specimen from Leticia have conspicuous dorsal patterns of dark brown blotches superimposed on a lighter ground color. Caqueta. In the broadened portion of the dark band above the urostyle there is a small. in Colomand from Trinidad. with a mean of 73.75 and a standard deviation of 5.) None of the type localities of the six separate names presently listed as synonyms are within Colombia. and foot are each crossed by two broad bands of dark brown which are coincident the legs are flexed. 44. and tibiatrix Lauren ti. elegant-looking frogs. seem to represent the variant described by Spix as H. . leg. p. will be referred to the genus Phrynohyas when they have been studied. the number of such teeth in three Trinidad specimens ranges from 51 to 56. The problem of variation in this species is no nearer Variation. dumb-bell shaped patch of the lighter dorsal ground color. Four other specimens. p. 1843.8 mm. 127) apparently was unaware that Commission (1958) placed the trivial name zonata Spix (as published in the combination Hyla zonata) on the Official Index of Rejected Names in Zoology and suppressed for purposes of priority both Acrodytes Fitzinger. These four specimens are not only smaller (60. all from La Providencia.S. 112) pointed out that there number is a distinct difference in the Leticia.3.6.32. A stripe of dark brown originates behind each eye and passes posteriorly around (but not through) the tympanum and terminates above the insertion of arm. A similar dark brown band crosses each forearm just above the wrist.

USNM 152248. a technique used by the Indians to maintain the rhythm of the strokes when paddling. San Juan de Arama. The pain kept me awake the first night. Tolima: Espinal. Villavicencio. . 400 m. It appeared toad-like but very alert. My hands had numerous scratches and small cuts and my left index finger a half-inch skinned place. and in spite of washing and wiping. CNHM . Thus. Rivero (1961. thickly wooded lot between houses in Leticia in the direction upriver from the basino. many stopped calling. Its vocal sacs were puffed out on the side of the neck and brown-gray in color. MLS CJG 263. think it —The skin secretions of this species are quite toxic and we pertinent to again quote Ross Allen's field notes that originally were published in Goin and Layne (1958. USNM 152198. MLS 258. turning its head from one side to another. Vaupes: Gino-Goje. Then I Its skin grabbed him as he started away and examined him more closely. USNM 152141. but in a few minutes I found one sitting on the edge of a rainwater pool. 265b. 1768) on the List of lished in the combination Specific Names in Zoology. it got worse. my hand so I put him in my frog sack. Morelia. natives along the Amazon call this the boatman frog in allusion to the fact that its voice resembles the tapping of paddles on the side of a canoe. Bolivar: Rio Vie jo. As has been pointed out by Lutz and Kloss (1952. all of which began to burn. Caqueta: La Provindencia. so after midnight I went to the place. Rio Manso. it produced . p. 656). p. 1824. but my hands then healed without a whitish mucus that covered further complications. upper Rio San Jorge. on the side was very granulated looking and while I was handling it. which was in a vacant. MLS 68. MCZ 15062. Los Micos. CNHM 61171. When I approached. Meta: Angostura No. State not known: Puerto Yaviya. I went back to camp and tried to wash off the poison and applied unguentine. the acid-like burn remained along with the swelling for about three days. The commission further ruled that the nominal species Rana venulosa Laurenti be interpreted by reference to the type specimen of Hyla zonata Spix. Remarks. The index finger and skinned place became fire-red. ANSP 25313." Specimens Examined COLOMBIA Amazonas: Leticia. UF 10021. the specific name of this well-known frog becomes officially and finally Phrynohyas venulosa. USNM 152140. C6rdoba: Catival. and venulosa Laurenti (as pubRana venulosa Laurenti. swollen. 81328. Rio Guayabero. 1843. p. 258a.—COCHRAN FROGS OF COLOMBIA AND GOIN 329 placed Phrynohyas Fitzinger. upper Rio Apaporis. 113): While standing guard over our precious bufeo [freshwater porpoise] I heard a chorus of frogs that was new to me. Norte de Santander: Astillero. USNM 152012-6. 131) points out that in Venezuela this species has the very appropriate common name "rana lechera. USNM 152047-50. 2298. . 2441 CJG (16). and puffed up.. 1.

P. the tongue extensively free behind.— Merrem. or if such spots are present they are much smaller than tympanum. hypochondrialis hypochondrialis to region of sacral hump Posterior surface of thigh plain or with a pattern but without conspicuous black and white markings. Suriname) . in most species (all but bicolor of the Colombian forms). Diagnosis. nicefori A distinct pair of white spots below the vent e P. 1802. p. 120. 113. l . 330 NATIONAL MUSEUM BULLETIN ECUADOR: El Oro. 1826. 1858. p. Key a 1 . p. 147.— Baudin. 1958. 1919.—Peters. Hyla bicolor.— Funkhouser. c 1 Upper surface of disks on fingers one and two without distinct white caps.. A distinct row of small white dots on each dorsolateral fold from behind No row .— MirandaRibeiro. boliviana of disks of fingers one and two with distinct white caps. the first toe longer than the second. Upper surface P. 1830. 1957. 3013-4. . 208.S. 220. 1830. p. 1868. Rana bicolor Boddaert).— Gorham. Posterior surface of thigh brightly patterned with black bl . Fitzinger. having the pupil vertical. bicolor Phyllomedusa bicolor (Boddaert) Plate 40a-c 1772. p. tympanum c2. 26.— B. p. Avila. p. 63. Phyllomedusa scleroderma Cope. 1961. —A large Phyllomedusa with the second toe longer than . p. Napo2507-8. rounded. white spots in the dorsal ground color along the sides between axilla and groin. 601. 1926.— Boulenger. blombergi eye to region of sacral hump of distinct white spots along dorsolateral region from behind eye P.U. 120. 3007-8. 1829. 1963.— Beebe. 40. 126. generally no dorsal pigment on tops of fingers.— Gunther.— 629. 427. fig. Montalvo. and in having very little or no webbing on the feet. these spots about size of c2.— Gravenhorst. 201 (type species. 70.. generally dorsal pigment on tops of fingers. EPN Genus Phyllomedusa Wagler Phyllomedusa Wagler. p. 1830.— Nieden. p. 1882a. 76 Pastaza. p. p. p. p.— Guerin-Meneville.—Shaw 1802. p.—Rivero. Pyllomedusa bicolor. Chicherota. b2 . 156. Calamita bicolor. — 1829. 26. 1950. km. 23.— Goin. P. and white markings. edentula No distinct pair of white spots below the vent e2 d l Some rather distinct.— Wagler. p. the palpebral membrane not reticulate. 112 (type locality. d 1 No series of rounded.— Spix. 42. p. Lutz. Dumeril and Bibron. to Colombian species of Phyllomedusa First toe longer than second. 1802. 1838. 170.—Tschudi. 201. p. 1799. p. 1799. 19 (type locality. — The frogs of this genus are characterized by Generic diagnosis. p. 1873a. 22. 103. 1923. 1803. P.— Goin and Layne. 337. 1820. p.. . 2 88 SUNHM EPN 2569. p. 1824. p. white spots in the dorsal ground color along the sides between axilla and groin. p. tarsius a2 . c1 .—Schneider. . southeast EPN of Buena vista. 3. Rana bicolor Boddaert. pi. 1958. 1841. p. and. Suriname). . 38. First toe shorter than second P. distinct. p.

Body rather elongate. tongue three-fourths as wide as mouth opening. Skin of upper parts glandular. head width. 33. not prominent. No patagium. — 73. low. Amazonas.2 mm. no vocal sac apparent externally. a distinct oval inner and a smaller but equally dictinct rounded outer metatarsal tubercle. Colombia... the upper jaw extending beyond lower. 43.. the upper lip not flaring out below it. when hind leg is adpressed. head length. Fingers unwebbed. no projecting rudiment of a pollex. third and fifth toes subequal.1 mm. — heel fails to reach the tympanum.4 mm. no dermal appendage on heel. no traces of a skinfold across chest. Skin of head not co-ossified with skull. interorbital distance twice the width of upper eyelid. 110. Snout short. reaching middle of disk of third which just covers the tympanic area. lying between the rather large. heels overlap slightly.FROGS OF COLOMBIA — COCHRAN AND GOIN 331 the first. 46. P. heel-to-toe. rather heavy series. their distance from tip of upper jaw about one-half that from eye. Tympanum moderately distinct. bicolor can be distinguished from all other species of Phyllomedusa now known from Colombia by having the second toe longer than the first. Toes with just a trace of web at base. . hand. 35.1 mm. tibia. calcarifer by the absence of web between the fingers. that of belly and lower surface of thigh uniformly granular.5 mm. Dimensions. disk of fourth toe covering about two-thirds the tympanic area. palpebral membrane not reticulate. posterior margin of tympanum bordered by what appears to be a broad low-lying process of the dorsolateral fold. It can be distinguished from Agalychnis spurrelli and A.. second toe longer than first. Vomerine teeth in two short. no tarsal ridge. ovate choanae. in postaxillary region distinctly narrower than greatest width of head. nostrils lateral. when hind legs are bent at right angles to body.7 mm.. roof of skull not exostosed. wide dorsolateral folds extending from behind eye to region of sacral hump. Eye rather small for a frog of this size. separated from eye by a distance about equal to one-fifth its own diameter. its posterior border free and unnotched. when limbs are laid along the side. no ulnar ridge. Description. skin of throat and chest fairly smooth. which is about equal to distance between nostrils. slightly projecting. knee and elbow fail to meet. fourth finger considerably longer than second. separated from each other by an interval equal to about twothirds their distance from eye. loreal region slightly concave and practically vertical. from Leticia.. about three-fifths the diameter of eye. truncate when viewed from above and truncate in profile. and with fingers and toes unwebbed except for a trace of web between bases of third and fourth toes. Head and body. its diameter equal to its distance from nostril. femur. 42. adult male. Can thus rostralis sharply defined. broadly pear-shaped. UF 10020.2 mm.

frogs of this species are used by the colors are separated line — — natives to induce vomiting. The row of white dots from eye to sacral hump will distinguish P. blombergi from all other Colombian species of Phyllomedusa at a glance. measuring 106. Santa Rosa de Sucumbio. Santa Rosa de Sucumbio) . tongue one-half as wide as mouth opening. Specimens Examined COLOMBIA Amazonas: Leticia. 39. p. that is practically identical to the described specimen except that the dorsal ground color has faded to a moderate bright blue and the subgular and pectoral ocelli are lacking. — Description. 23. round. NATIONAL MUSEUM BULLETIN 2 88 — Very dark gimmetal blue dorsally. 1963. The above the vent coincides with the lines on the heels when the legs are flexed. black-bordered. elongate. of forearms. On the lateral surfaces of forearms.S. an adult male. Under the chin there is a small 8-shaped white spot with a black border. yellowish white. Goin and Layne. PutuGorham. On all where the dorsal and ventral ground colors meet (except just above the vent and on the lateral surfaces of the forearms and hands and feet) the two ground colors are separated by a very conspicuous row of rounded black-bordered ocelli-like white spots. and under chin gray Color in alcohol. The other specimen at hand (CNHM 81929) is an Variation. p. CNHM 81929. hands. 41 (type locality. and feet and above the vent the two ground places by a distinct. Putamayo. As has been pointed out (Funkhouser. white spots below the vent.332 U. Phyllomedusa blombergi Funkhouser. and under lateral portions throat. from Rio Rumiyacu. cadaverish under the feet. black-bordered white markings. 1958. free and . although there is a small ocellus under the base of each arm. and on the chest there is a single round white spot with a black border. A small Phyllomedusa with a truncate snout and welldeveloped parotoid glands and a distinct row of small white dots extending from upper eyelid along dorsolateral fold to sacral hump. Colombia. UF 10020. — — Diagnosis. mayo. Rio Rumiyacu. Vomerine teeth in two tiny patches between the somewhat squarish moderate-sized choanae. There is a pair of very distinct.5 millimeters in head-body length. The voice of frogs of this species is a loud nasal snarl. Remarks. 113). SUNHM 13241 (holotype). 1957. The lower lip is margined in part with similar. Colombia. Phyllomedusa blombergi Funkhouser Plate 40d-f 1957. rest of ventral surfaces a dirty. p. elongate. p. black-bordered white line.

truncate in profile. Possibly a juvenile. fifth a disk longer than third.8 mm. Head and body. 9 mm. Very distinct white lines bordering outer margin of hand and ulnar fold and outer margin of foot and tarsal fold. 14. 34. very slightly projecting. their distance from end of snout about one-half that from eye. nostrils entirely lateral. the upper jaw not extending considerably beyond lower. when hind leg is adpressed. heels overlap appreciably. brownish gray below. Canthus rostralis very sharply defined. heel-to-toe. no vocal sac apparent. which is about equal to its distance between nostrils. skin of throat and chest smooth. separated from each other by an interval nearly equal to their distance from eye. a weakly developed ulnar ridge.. Body somewhat elongate. this line just matching white line on heels when legs are flexed.. tibia. when hind legs are bent at right angles to body. loreal region flat and practically vertical. Skin of head not co-ossified with skull. disk of fourth covering about onefourth the tympanic area. Fingers unwebbed. Skin of upper parts fairly smooth except for dorsolateral folds that extend from posterior portion of eye to sacral hump and bear a distinct but narrow row of small. interorbital distance greater than width of upper eyelid.FROGS OF COLOMBIA —COCHRAN AND GOIN 333 very shallowly notched behind. Color in alcohol. Eye moderate but prominent. that of belly and lower surface of thigh uniformly and very finely granular. palpebral membrane not reticulate. a distinct oval inner and a smaller indistinct rounded outer metatarsal tubercle. in postaxillary region distinctly narrower than greatest width of head. Snout moderate. 11. Upper and lower lips margined with a distinct white line. about one-half the diameter of eye. No patagium. Toes unwebbed. no dermal appendage on heel. reaching to middle of disk of third which covers about onethird the tympanic area. 19. its diameter one-third again as great as its distance from nostril. fourth finger longer than second. hand. A very distinct short horizontal white line above vent. a rather narrow glandular ridge encircling upper part of tympanum and extending downward toward shoulder. tarsal ridge present. a very distinct row of small white dots extending from upper eyelid along dorsolateral fold to sacral hump. first toe longer than second. Tympanum very distinct. rounded pustules.. no traces of a skinfold across chest.7 mm.9 mm. 13.3 mm. when limbs are laid along the side.. knee and elbow overlap considerably. head width.. heel reaches middle of eye. pointed when viewed from above. each pigmented a creamy white. Dimensions. Slate blue above. sex not determinable. femur. no projecting rudiment of a pollex. separated from orbit by a distance about equal to onethird its own diameter. A rather indistinct white line on each side extending posteriorly from angle of jaw to about middle of — — . roof of skull not exostosed.7 mm. 13.2 mm. the upper lip not flaring out below it.. head length.

— Gans. p. Bradymedusa hypochondrialis. 1926. 1841. from Villavicencio. On — Phyllomedusa hypochondrialis hypochondrialis (Dawlin) Figure 39 1803. 1896. p. broadly triangular when viewed from above. P. tongue three-fifths as wide as mouth opening.334 U. NATIONAL MUSEUM BULLETIN 2 88 each side posterior to this line are three rounded white dots anterior to groin.— Gravenhorst. 1963. 204— Mehely. p. 1820.— Nieden. rounded. and well separated.— Bles. 1961. 1901. 112. A narrow margin of white on lateral side of little finger to base of disk. A pair of rounded white dots are in middle of throat region and a single ocellus-like white spot is in middle of pectoral region. — —A small. p. truncate in profile. Cope. 44. its posterior border free and unnotched. 24. 1919. 1829. From blombergi it can be distinguished by the lack of a row of white dots along each dorsolateral fold from behind the eye to the region of the sacral hump. 1868. Colombia. 1923. 355. Rivero.— Stebbins and Hendrickson. Miranda-Ribeiro.— B. the upper — CNHM . 29. Description. p. p. 303 Gorham. 1820.— Berg. p. 150. p. 1826. where it stops abruptly. 1868. 430. 255. p. 81771. — Phyllomedusa bicolor. 264.S. slightly projecting. p. 230. p. Snout short. On the hind limbs the dorsal pigment covers dorsal surface of fifth toe and half of dorsal surface of fourth toe. — Giinther. Suriname). On the forelimbs the dorsal pigment extends to base of fingers. Phrynomedusa hypochondrialis. Merrem. 63. 1960. 1950. p. Phyllomedusa hypochondrialis hypochondrialis. we know nothing of this species' geographic range or variation. 1923a. 1862. 1858. — Fitzinger. 120 (part). Vomerine teeth absent. Dumeril and Bibron. 605 Musshoff. 601. p. broadly pear-shaped. 213. 105. choanae small. lS62b. — p. their distance from end of snout about one-half that from eye. p. Since the type is the only known specimen. — brightly colored Phyllomedusa with the first toe longer than the second. Canthus rostralis well defined. 1899. 313. — — — — Cope. 1904. 1961. 1939. Meta. 1906. Diagnosis.— Gadow. 1882a. p. 1957 p. 1882b. p. 208. p. 170.— Funkhouser. p.— Miranda-Ribeiro. Remarks. — Boulenger. p. Pithecopus hypochondrialis. p. Calamita hypochondrialis. 1923. 629 (part). Lutz. p. 29 (type locality.— Budgett. loreal region nearly flat and very nearly vertical. side. hypochondrialis hypochondrialis can be distinguished from all Colombian Phyllomedusa except blombergi by the combination of a bright black and white pattern on the rear of the thigh together with the first toe being longer than the second. Phyllomedusa hypochondrialis. 527. 337 Lutz and Lutz. nostrils lateral.— Beebe. p. 1959. separated from each other by an interval slightly greater than their distance from eye. 3. Hyla hypochondrialis Daudin. 1926. 1905. p. — p. 219. the upper jaw extending hardly at all beyond lower. p. sloping but very slightly.

separated from eye by a distance equal to about one-fourth its own diameter. when hind leg is adpressed. disk of fourth covering about two-thirds the tympanic area. in postaxillary region a little narrower than greatest width of head. heel reaches middle of eye. fifth longer than third. disk of third covers about three-fourths the tympanic area. first toe longer than second. no true tarsal ridge. palpebral membrane not reticulate. its diameter half again as great as its distance from nostril. Body not elongate for a Phyllomedvsa.— FROGS OF COLOMBIA —COCHRAN AND GOIN 335 lip flaring out hardly at all below it. a distinct oval inner and a smaller rounded outer metatarsal tubercle. no dermal appendage on heel. Phyllomedusa hypochondrialis hypochondrialis. interorbital distance slightly greater than width of upper eyelid. a weak ulnar ridge. about one-third the diameter of eye. USNM 15249 (X 2). . when limbs are laid Figure 39. Fingers not webbed. which is about the distance between nostrils. no projecting rudiment of a pollex. Eye small. not prominent. fourth finger nearly as long as third. Tympanum distinct. Toes without webs.

The light margin with the contact in come to as a light lateral stripe that passes above posteriorly continues of lip arm and continues along the side half-way to the groin. and upper arms are conspicuously marked with dark curlicues. A light stripe on the lateral margin of each hand extends along the ulnar ridge to elbow. — A dark purplish-blue above. this margin broadening below the eye with margined Upper lower margin of the eye. curlicue-like. 97090) from Pernambuco.' —It not at is all surprising to find this species in the parts Colombia east of the Andes.S.. dark. hypochondrialis azurea Cope from southern South America. adult female. chest. A narrow glandular fold passes downward behind the tympanum. The dorsal surfaces of the thighs are faintly pigmented. head length. along the lateral margin of each tarsal region to and across the heel. MLS USNM SURINAME: 245. 11. CJG 2318-9. no vocal sac. We have seen a cotype (TJSNM 5832) from Paraguay and two specimens (USNM 48857.8 mm. lip but posterior to this they become distinct. heels overlap considerably. The anterior and posterior surfaces of the thighs are pale with distinct. Onverwacht. Skin of head very thick but not co-ossified with skull. white. knee and elbow overlap slightly.2 mm. pale yellow below.. Villavicencio. tibia. skin of throat. more or less vertical bands. 152249-60. Skin of upper parts rather smooth. 12. 14. and for the time being are willing to go along with Rivero that azurea of is a recognizable race of P. CJG MLS 2422. and lower surface of thigh uniformly and very finely granular. 51) recorded it from the Venezuelan Guayana. Brazil. feet. 16. p. Remarks. Head and body.NATIONAL MUSEUM BULLETIN U. head width. 245a-b.. — Dimensions. no traces of a skinfold across chest. No patagium. Below this lateral light stripe there are faint vermiculations for" about half-way to Color in alcohol. 35 mm.. femur. hypochondrialis. hand.3 mm.. roof of skull not exostosed. heel-to-toe. belly. A similar stripe on the lateral margin of each foot extends posterior surfaces of the thighs. 23 9 mm. mm. 151. about four on each thigh. dark. thus giving an impression of a tarsal ridge. no discernible parotoid glands. A short horizontal white stripe lies just above the vent. 81771-4. dorsal band of pigment separating the dark stripes on the anterior and The concealed surfaces of shanks. this groin.. CNHM . Rivero (1961. of what is called P. Specimens Examined COLOMBIA Meta: Acacias. markings. 336 2 88 along the side.4 mm. when hind legs are bent at right angles to body.

Fingers entirely free. on the disks of fingers one and two. pointed when viewed from above. p. not projecting. separated from each other by an interval equal to about four-fifths their distance from eye. Description. p. From tarsius it differs in lacking white caps Diagnosis. the upper jaw extending but slightly beyond lower.third than its distance from nostril. rounded white spots along the side anterior to the groin. — Funkhouser. Colombia. the upper lip flaring out strongly below it. Among the Phyllomedusa now known from Colombia. about one-half the diameter of eye. heel reaches posterior margin of eye. . oval series lying close together and between the rather large. separated from eye by a distance about — equal to one-fourth its own diameter. B. in postaxillary region distinctly narrower than greatest width of head. Loveridge. if such spots are present. very prominent.FROGS OF COLOMBIA —COCHRAN AND GOIN 337 Phyllomedusa nicefori Barbour Plate 42d-f 1926. Meta. Gorham. slightly greater than distance between nostrils. and without distinct white caps on the upper surfaces of disks of fingers one and two. nostrils lateral. Tympanum very distinct. 1957. just reaching to disk of third which covers about three-fourths the tympanic area. they are much smaller than in boliviana. p. 1929. interorbital distance about V/i times the width of upper eyelid. fifth distinctly longer than third. the second. first toe longer than second. their distance from end of snout about twothirds that from eye. Canthus rostralis well defined. finger considerably longer . p. boliviana. From boliviana it differs in lacking rounded white spots anterior to the groin or. distinct. tongue one-half as wide as mouth opening. 47 (as niceforoi). MCZ 11611 (holotype). its posterior border free and truncate but not notched. loreal region concave and oblique. when hind leg is adpressed. — — 1963. and tarsius by the lack of conspicuous black and white markings on the rear surface of the thigh. Eye large. Lutz. Snout large. Villavicencio) 1950. —A moderate-sized Phyllomedusa with the first toe longer than. its diameter greater by one. Body elongate. sloping in profile. no tarsal ridge. without conspicuous. Meta. no rudiment of a pollex. disk of fourth covering about three-fourths the tympanic area. 24. from Villavicencio. Phyllomedusa nicefori Barbour. Vomerine teeth in two short. —Barbour and 602. ovate choanae. toes without webs. p. fourth than second. 191 (type locality. Colombia. this species may be distinguished from all except edentula. From edentula it differs in having a pair of rounded white spots below the vent. an oval inner but no discernible outer metatarsal tubercle. which is relatively wide. 319. no|dermal appendage on heel.

33. Boulenger. rounded. Dimensions. 48) was so unwilling to commit herself concerning the status of this form as there is a perfectly good type available. indicate that this species is rather constant in its characters. The pair of white spots below the vent is conspicuous in all of them.. vocal sac not apparent.3 mm. Slightly mottled on sides of body and limbs where the dorsal and ventral colors meet. USNM 118171. belly..— Head and body. 395 (type locality. a pair on base of each leg near the vent. Diagnosis. head length. and lower femur uniformly granular. Phyllomedusa boliviano. Parotiod glands present but not pronounced. 47. femur.9 head width.. CNHM 63860.— Funkhouser.. The series of specimens at hand. Remarks. 341. 1963.— Gorham. We are unable to understand why Funkhouser (1957. No patagium. Phyllomedusa boliviano. 57. p. The warts on chest and near the vent stand out as clear yellowish marks.. USNM 157261-2. MZUM 54634.5 mm. 35. foot. The dorsal and ventral ground colors come together along the sides anterior to the groin with a slight mottling effect but without any development of clear. Norte de Santander. when hind legs are bent at right angles to body.S. 1957. CNHM 81769-70. 1923. mm. extending posteriorly nearly to sacrum.9 tibia. 602. a pair of yellowish warts in tandem on pectoral region of chest. mm. 1950. heels overlap greatly. Boulenger Plate 40g-i 1902. skin of throat.6 mm.. Chulu- mani). 76. moderate-sized Phyllomedusa with well-developed . 21. p.338 NATIONAL MUSEUM BULLETIN U. p. — Uniform very dark (midnight) blue above. — Specimens Examined COLOMBIA Antioquia: Porcecito. 2 88 when limbs are laid along the side. Bolivia. p. —A p. a short series behind the angle of each jaw and well-developed patches of them along sides of body. dirty gray below. pp. Santander: Aratoka. 25.8 mm.2 mm.—B. No data: MLS 244. white spots the size of the tympanum. and a series of five on ventral surface of base of arms. Color in alcohol. Skin of upper parts glandular but not warty.— Nieden. San Calixto. including a number of rather fresh topotypes. knee and elbow overlap distinctly. hand. 23. 49. 25. Lutz. chest. MLS (a). Meta: Villavicencio. Two of the specimens (CNHM 81769-70) are more reddish than the others but this may be due to preservative. no trace of a skinfold across chest. MCZ 11611 (type).

their distance from end of snout about one-half that from eye. a rather narrow glandular ridge passes down and behind the tympanum. fourth finger considerably longer than second. two short series. palpebral membrane not reticulate. not at all superior. a weak ulnar ridge. separated from eye by a distance equal to one-fourth its own diameter. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. when limbs are laid along the side. no vocal sac. squarish choanae. lying close together between the small. Vomerine teeth 62. the Colombian species with which it is most likely to be confused. can be distinguished by the white spots along the side. fifth longer than third. no projecting rudiment of a pollex. in postaxillary region a little narrower than greatest width of head. From P. about one-third the diameter of eye. inconspicuous parotoid glands extending from the eye to region of the sacral hump. in when hind legs are bent at right angles to body. broadly triangular when viewed from above. white spots in the dorsal ground color along the sides parotoid glands. roof of skull not exostosed. disk of fourth covering about one-half the tympanic area. boliviano. Eye not large nor particularly prominent. no dermal appendage on heel. In nicejori there may be a few white spots in this region but if so they are small and granular rather than large (the size of the tympanum) and rounded —MLS as they are in boliviano. disk of third just covers the tympanic area. p. from La Providencia. No and rather coarsely granular. Toes without webs.FROGS OF COLOMBIA — COCHRAN AND GOIN 339 first toe longer than second. no traces of a skinfold across chest. skin of throat and chest very finely granular. Skin of upper parts very finely glandular. nostrils lateral. nearly as long as third. Skin of head not co-ossified with skull.. slightly projecting. Body rather elongate. a pair of broad. Caqueta. between axilla and groin. the upper lip flaring out but slightly below it. first toe longer than second. broad. which is relatively wide and slightly greater than distance between nostrils. knee and elbow overlap Description. . truncate and slightly sloping in profile. its posterior border extensively free and unnotched. adult female. Colombia. and some rather rounded. Fingers not webbed. Canthus rostralis well defined. that of belly and lower surface of thigh uniformly considerably. heels patagium. Snout short. Tympanum distinct. its diameter slightly greater than its distance from nostril. when hind leg is adpressed. heel reaches anterior margin of eye. nicefori. overlap greatly. interorbital distance slightly greater than width of upper eyelid. the upper jaw not extending appreciably beyond lower. tongue three-fourths as wide as mouth opening. separated from each other by an interval equal to their distance from eye. a very weak tarsal ridge. the clear. low. loreal region slightly concave and nearly vertical.

bicolor in having the first toe longer than second and in lacking brilliant markings on the legs. 24.49-50 [1947. heel-to-toe. 27 mm. of Rio Napo). BM 1901.. BOLIVIA: Chulumani. and a pair of white spots be- second. MLS 2.4 Color in alcohol.6 mm. 603. USNM Phyllomedusa tarsius (Cope) Plate 41 1868. mm. cadaverous gray below. 113 (type locality. and on the concealed surfaces of shanks and feet.2. head length. — Specimens Examined COLOMBIA Caqueta: La Providencia. The two topotypes of boliviana in the British Museum differ from our specimen in that they have a distinct pair of white spots below the vent while the specimen from La Providencia has only one pale spot on one side below the vent. 340 U. Distinct whitish rounded spots along the sides between axilla and groin. Boulenger. the sides and thighs marked with pale fleckings. 1922. in his opinion it was closer to boliviana. 343. 1923.000 m. 62. Remarks.8. white caps on disks of first above the vent and on base of low the vent.. NATIONAL MUSEUM BULLETIN 2 88 — Dimensions.jeltoni so we sent it off at once for our friend Benjamin Shreve to examine. edentula. 32. 44. Whitish blotches on the lower surface of base of each arm.. Ruthven. Upper mouth 1882.—B. —Blue-black above.8 mm. mm.— . 1963. and a rounded distinct light spot in the middle of chest. an identification with which we concur. on the concealed surfaces of thigh. a white line each heel.— Funkhouser. p. femur. p. Margin white line extending from posterior corner of lip to above arm. Head and body. He reported that while it was close to feltoni. Pithecopus tarsius Cope.32-3] (cotypes) 118699-700. —A large Phyllomedusa with the first toe longer than the and second fingers.. Peru. 51. Buena vista (Department of 62.4 mm. p. Nieden.— Gorham. 428. Lutz. The ulnar and tarsal ridges lined of lower lip white. it differs in having white caps on the disks of first and second fingers. 21 mm. Whether this difference has any significance we cannot say at this time. with less-distinct ones on the upper arm. Diagnosis. 1957.2. Santa Cruz). 57. p. 21. 1950. and paired white spots below the vent. tibia. a with white. A light line above the vent and similar horizontal light lines on each heel. — Amazon below — Phyllomedusa tarsius. From P. a white line above the vent and on each heel. head width. p. 1882a..S.22. hand. This species differs from P.1 20.. to which it is perhaps most closely related. p. . on anterior and posterior surfaces of thigh. When we first received the single Colombian specimen we thought it might be P. p.

its diameter one. interorbital distance slightly greater than width of upper eyelid. no vocal sac.5 mm. fifth longer than third. 3.15.— 1902.1 mm. 42. Body rather elongate. no true ulnar ridge but side of hand and arm marked by a white line which gives the impression of a ridge. tibia. no inguinal gland. Skin of throat very finely granular.. a distinct oval inner and a rather obscure outer metatarsal tubercle. the upper jaw extending very slightly beyond lower. not particularly prominent. and above 337-262—70 23 . Snout moderate.third again as great as its distance from nostril.5. head length.. Skin between axilla and groin somewhat granular and wrinkled. about three-fifths the diameter of eye and nearly in contact with it. fourth finger considerably longer than second and practically as long as third. a fairly well-developed parotoid gland extending from upper eyelid to sacrum. spots superimposed on some of the granular elevations. its posterior border free and shallowly notched.5 mm. reaching nearly to disk of fourth which covers about one-half the tympanic area. Head and body. Colombia. their distance from end of snout about one-half that from eye. Vomerine teeth in two short.FROGS OF COLOMBIA — COCHRAN AND GOIN 341 BM Description. female. knee and elbow overlap appreciably when hind legs are bent at right angles to body.3 mm. first toe longer than second. Toes unwebbed.500 feet. roof of skull not exostosed. that of belly and lower thigh uniformly granular. which is obscured by parotoid gland. slightly projecting. — Dark blue above. No patagium. heels overlap appreciably. 95.. 32. no definitive tarsal ridge but a pale line along outer surface of foot and tarsus which gives the impression of a ridge. slightly curving. hand 28. separated from each other by an interval about equal to their distance from eye. tongue two-thirds as wide as mouth opening. Skin of upper parts very finely glandular. no projecting rudiment of a pollex. Skin of head not co-ossified with skull. disk of third covers about three-fourths the tympanic area.. somewhat pointed when viewed from above. an elongated heart-shape. the upper lip flaring out slightly below it. Can thus rostralis well defined. head width. in postaxillary region a little narrower than greatest width of head when hind leg is adpressed. a dirty bluish brown below. Fingers unwebbed.4 — mm. heel reaches anterior margin of eye when limbs are laid along the side. no dermal appendage on heel. 28. sloping in profile. Dimensions. rather rounded series lying between the moderate-sized oblong choanae. traces of a skinfold across chest.. Eye moderate.9 mm. Tympanum distinct.25. Cundinamarca. 63. On the side the dorsal and ventral colors are separated by some white Color in alcohol. foot.. Palpebral membrane not reticulate but with some pigment. nostrils more lateral than superior. from Villeta. loreal region slightly concave and nearly vertical. 45. chest fairly smooth.5 mm. femur.

5 millimeters.5. a single pectoral white spot.25. p. and this pigmentation extends down a millimeter or two on the membrane itself. p. p.5. Rio Pastaza watershed).S. lower lip white. either on the thighs or on the sides. somewhat triangular in shape. Both anterior and posterior faces of thighs are flecked with whitish lavender spots like those along the sides. —The Colombian specimen very similar to virtual topotypes from Peru.9. — — — A large Phyllomedusa with pigment on tops of the two Diagnosis. 1963. Chanchamayo. an indistinct white line along the outer side of hand and lower arm and outer side of foot and tarsus.6-7.15. and without two rounded white spots below the vent. Gorham. tongue four-fifths as wide as mouth opening. Funkhouser. Phyllomedusa edentula Andersson. a female from Pebas. nicejori.. horizontally oriented choanae. The margin of palpebral membrane is purple. Lower eyelid white. Variation. while in a specimen from Pebas the ulnar and tarsal stripes are not so pronounced although they are well developed in the other specimen from the same locality. 3. The longer first toe serves to differentiate P. Phyllomedusa edentula Andersson Plate 42a-c 1945. BM 67. Snout moderate. is Specimens Examined COLOMBIA Cundinamarca: PERU: Pebas. 74906.5. Dorsal ground color is present on upper arm and extends to disks of third and fourth fingers.342 U. Description. Villeta. Choc6. and the lack of the two white spots below the vent distinguishes it from P. edentula from P. In one of the Peruvian specimens (BM 1908. which it approaches in general habitus and pattern.29. 1908.17. It is present on thigh and extends to disks of fourth and fifth toes. which it approaches in size.45) the flecks are not so well developed. its posterior border extensively free and unnotched. A pair of white spots below vent and a couple of indistinct white spots on lower surindistinct white line behind angle of face of each thigh. —CNHM . BM 1902. 84 (type locality. 23. NATIONAL MUSEUM BULLETIN 288 these are rather numerous whitish-lavender flecks on the smoother skin above granular region. Vomerine teeth in two small patches lying not too close together between the large. disks of thumb and first finger white.500 BM ft. The largest specimen. Colombia. jaw on each side. 1957. from Sautata. has a head-and-body length of 110. a short. Ecuador. with the first toe longer than second. a white blotch at pectoral portion of base of each arm.29. outer fingers. oblong. A white line across each heel and another above vent. bicolor.45. 43.

A low but distinct glandular dorsolateral fold extends on each side from behind eye to region of sacral hump. all gray. Head and body. another row along ulnar region. scattered white dots along the side between the dorsal ground color and the ventral ground color. hand. when hind legs are bent at right angles to body. about three-fifths the diameter of eye. when limbs are laid along the side. no projecting rudiment of a pollex. A distinct.5 mm. somewhat projecting. no trace of a skinfold across chest. Dimensions. heel reaches anterior border of tympanum. white.. much longer than third.8 co-ossified — mm. the first toe longer than second. Fingers unw ebbed. separated from each other by an interval about equal to their distance from eye. interorbital greater than width of upper eyelid. no dermal appendage on heel. which is cadaverish Color in alcohol. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle.. rounded. out but very slightly below it. No patagium. — Gunmetal blue on exposed dorsal surfaces except top of first two fingers and first three toes. that of belly and lower its surface of thigh uniformly and more coarsely granular. their distance from tip of snout about equal to their distance from eye. tibia. 91. truncate in the upper jaw extending but slightly beyond lower. 60. its diameter half again as great as its distance from nostrils. Skin of head not with skull. heel-to-toe. the upper profile.FROGS OF COLOMBIA —COCHRAN AND GOIN 343 more pointed than rounded when viewed from above. lip flaring membrane not palpebral reticulate. fourth finger considerably longer than second. in postaxillary region distinctly narrower than greatest width of head. heels overlap greatly... when hind leg is adpressed. ocellus-like pectoral spot but no white spots below vent. 31. separated from eye by a distance equal to one-half is own diameter. of Few somber-looking Phyllomedusa.. reaching to middle of disk of third which covers about two-thirds the tympanic area.4 mm. head width. —We the species America. are not satisfied with the present arrangement of somber-colored Phyllomedusa from western South of these species have been seen in life by other than . knee and elbow overlap slightly. Eye moderate. nostrils lateral.5 mm. disk of fourth covering about one-half the tympanic area. Canthus rostralis well defined. Small.. no tarsal ridge. Skin of throat and chest finely granular. 37. which distance slightly about equal to distance between nostrils. 30 mm. no vocal sac. Anterior and posterior concealed surfaces of the thigh lavender. adult female. ulnar ridge a row of very low.5 mm. head length. loreal region somewhat concave and nearly vertical. Tympanum very distinct. femur. Body elongate. Toes unwebbed. 42 mm. roof of skull not exostosed. 29. All-in-all a rather Remarks. wart-like structures. not particularly prominent. A row of white dots from posterior angle of jaw to above axilla.

distinct white line. Specimens Examined COLOMBIA Choc6: Sautata. nicefori and to P. NATIONAL MUSEUM BULLETIN 2 88 the collectors. but she differs from that species in her much smaller size (62. This specimen has been directly compared with the type of edentula and differs from it in the same respects.S. and no one person has collected many examples of the named species. if in fact the latter two are really distinct. NR collection. edentula. in head-body length) than the two Colombian specimens. the recent monographer of the group saw type material of only four.6 mm. although the type is slightly smaller (83. The two Choco specimen were in amplexus when taken. So far we have seen type material of eleven and are not yet ready to assign more than relatively few of the names with any degree of confidence. but for the present we see no alternative. This frog seems most like P. Colombia. The two specimens for which we are using the name P. cannot be assigned to any species we now know. In these the female has teeth in both ridges but the male has them in the left ridge only. The two specimens here called edentula have been directly compared with the type of that species and the three seem to be conspecific. It is true that the type was described as toothless. .5 as compared to 91. ECUADOR: CNHM 74906-7. orcesi. Rfo Pastaza watershed (type). Phyllomedusa species Figure 40 A single specimen of Phyllomedusa from Serrania de La Macarena in Meta. Of the 17 species of phyllomedusids described from the northern half of western South America. edentula are certainly closely related to P. and in the fact that along her sides the dorsal and ventral ground colors are separated by a nearly complete.5 millimeters in head-body length.7 mm. in having low but distinct triangular dermal appendages on the heels.5 and 96. but there are two low ridges of the same size and position as the tooth ridges in the two specimens at hand. It somewhat disturbs our sense of zoogeography to assign specimens that were collected practically on the Darien border of Colombia to a species described from the eastern side of Ecuador. for our two Colombian specimens of edentula).344 U. The specimens of edentula differ most conspicuously from those of the latter two species in their larger size and in lacking two light spots below the vent. On both sides this line is interrupted in the region of the groin to form a short row of elongated pale spots. This individual (CNHM 81338) is an adult female measuring 62.

40. 300 m. —A pair of postorbital projections = Hyla baudin} on the fronto- depressor mandibulae with two distinct heads.) Since on the basis of the single specimen at hand we cannot tell whether this example of a Phyllomedusa simply represents a local race of edentula or whether it in fact represents a distinct species. parietal bones. we refrain from proposing a name for it at this time. M. There is some evidence of endemism in Serrania de La Macarena. Genus Smilisca Cope 1865.. Smilisca daulinia Cope Dum6ril and Bibron). 194 (type species. which is an isolated mountain range. Specimens Examined COLOMBIA Meta: Serrania de La Macarena. AND COIN CNHM 81338 (same 345 size). .— FROGS OF COLOMBIA Figure —COCHRAN Phyllomedusa species. described as a new species in this volume. Generic diagnosis. also comes from there. p. (Gastrotheca medemi. Smilisca Cope. Rio Guapaya. CNHM 81338.

S. from Rio Calima above Cordoba. vocal sac in male subgular with a tendency toward pairing.— Werner. 80. 4.— Nieden. by the above two characters. Snout short. p. 261. Fingers . 416. Smilisca p haeo la. its posterior border slightly free and very shallowly notched. p.— 618. transverse series lying close together on a level with the posterior borders of the small. Heatwole. and Stuart. 1946. 358 (type locality. p. Peters. p.— Taylor. 1901. p. its diameter equal to its distance from nostril. 271. p. Colombian Species of Smilisca dark stripe. Diagnosis. 22.— Giinther. broadly oval. p. 1897. Turbo). 589. NATIONAL MUSEUM BULLETIN 2 88 one of which originates on the squamosal.— Smith and Taylor. Eye moderate.— Gaige.— Liu. p. 1923. of tympanum eye two-thirds or more the diameter S. 402. tongue two-thirds as wide as mouth opening. Canthus rostralis well denned. Key a 1 . 88.— Taylor and Smith. gabbi. 1862.— Taylor. for gabbi lacks the postocular dark stripe and has a tympanum not over one-half the diameter of eye. A distinct postocular No . Colombia. not particularly prominent. p. palpebral membrane not reticulate. phaeota may be distinguished from S. p.— Breder. the only other species of Smilisca known from Colombia.346 U. 1862b. p. Hyla phaeota Cope. 300. 1937. 1964. somewhat oval choanae. p. rather rounded when viewed from above. —Starrett. which is relatively wide and slightly greater than distance between nostrils.— Gorham. separated from each other by an interval nearly equal to their distance from eye. Vomerine teeth in two short. 11 (type locality.— Dunn. phaeota hardly more than half the diameter S. 15. stripe that includes the — Description. interorbital distance about equal to width of upper eyelid. 1948. Hartweg. Rio Esnape.S. the upper lip flaring out rather strongly below it. their distance from end of snout about one-half that from eye. 1935. 1960. loreal region concave and somewhat oblique. 1945. Hyla baudini dolomedes Barbour. Colombia. p. 413. gabbi Smilisca phaeota (Cope) Plate 43d-f 1923. not projecting. 1963. —A moderate-sized Smilisca with a postocular dark tympanum and with the tympanum at least two-thirds the diameter of eye. Valle. tympanum eye of a2 to postocular dark stripe. about the diameter of eye. rounded in profile.— Boulenger. 837. separated from eye by a distance nearly equal to one-third its own diameter. 1942. — Sambu Valley). p. and Knight. Sexton. 1882a. 34. CJG 2252. p. 269. 1961. the upper jaw extending somewhat beyond lower. Panama. 1960a. p. p. 215.— Goin. Tympanum very distinct. 1931. 1873. p. nostrils nearly lateral. p. 1952.

head width. reach- ing to middle of disk of third which covers about one-third the tym- panic area..FROGS OF COLOMBIA webbed — COCHRAN AND GOIN 347 at base. heel-to-toe. 55.8 mm. Variation. in postaxillary region a little nar- rower than greatest width of head. top of thigh pale gray with pepper-like dark flecks. and terminates above shoulder. light margin.. the pattern is very much the same on millimeters. knee and elbow meet snugly.6 mm.2 mm. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. head length. Skin of upper parts smooth.2. Also. In some. top of shank pale gray with several indistinct dark gray crossbands. adult male.. third and fifth toes subequal.3. heels overlap greatly. 16.6 mm. hand. when hind leg is adpressed. heel when limbs are laid along the side. A rather distinct pattern of dorsal dark blotches and bands across the tops of the thighs and shanks evident in most of the specimens. Toes more than one-half webbed. 20. roof of skull not exostosed. there is a light line along the entire edge of the upper lip. 25. In all speci- is mens a light-bordered. 19. —Above. vocal sac well developed. no ulnar ridge. In the fresh material at hand... when hind legs are bent at right angles to body. all three largest males at specimens. Color in alcohol. no projecting rudiment of a pollex. dark. phaeota the females are distinctly larger males. one (CJG 2250) 71.3 hand (CJG 2251-53) measure 54.8 mm. a rather narrow glandular ridge encircling upper part of tympanum. postocular stripe that includes the tym- panum is conspicuous. — Dimensions. Skin of head not coossified with skull. no dermal appendage on heel.2 S. femur. the dark blotches on the back and the . —In light gray (CNHM 54768) millimeters in head-and-body length and the other The than the measures 69. Ventrally the specimen is very pale cadaverish gray except under chin and throat where it is rather heavily flecked with fuscous.6 millimeters in head-body length. Loreal region a little darker than top of head. with mottlings on the dorsum of dark gray. disk of fourth toe covering about one-fourth to one-third the tympanic area. 40. Of two large females at hand.. and 55. a very distinct dark postocular stripe runs posteriorly from the hind corner of eye. pale. Head and body. In fresh material the dorsal blotches are dark gray. skin of throat and chest finely granular. a distinct skinfold across chest. includes all of tympanum. 55. that of belly and lower surface of thigh uniformly and more coarsely granular. fourth finger considerably longer than second. no tarsal ridge. Body moderately elongate. tibia. the web on fourth toe reaching slightly distal end of antepenultimate phalanx. This dark stripe is bordered below by a very narrow. differing only in intensity. 33 mm.6 mm. on a light gray dorsal ground color. No patagium. reaches tip of snout.

82. with dark mottlings on the back. Chigorodo. Pichincha Province: vicinity of Santo Domingo de los Colorados. 151883-4. CJG 2463 (11). -Although the type is from Colombia.3.1 to 23. NATIONAL MUSEUM BULLETIN dark crossbands on the 2 88 dark For example.12. Remarks. CNHM BM CNHM AMNH CNHM CNHM CNHM BM 4451-3. Rio Rosario. some long-preserved specimens are now simply dark brown. Four specimens (CJG 2310-13) were collected east of Tumaco. CJG 2310-3. doba. p. 4055-6. 2964. Peiia Lisa. Imbabura Province: V/i km. JAP 2916-8. 81857. near Cachabi. 2829-30. Golfo de Uraba. ground a palm The in These specimens were collected on the ground. Playa de Oro. 18 km. 63881. 1882a. absent. 103 (type locality.69-70. 4124-5. UV 346-7. p. JAP 4014. Rio Raposo. leg stand out prominently.16. Rio San Juan. 1913. Rio Calima. 1913. the dorsal markings are very dim or. CJG 2252.11. BM USNM USNM BM Smilisca gabbi (Cope) Plate 43a-c 1876. Caldas: Pueblorrico-Santa Ceteilia. near Sipurio. USNM 152153-5. JAP 2814-24. and that night one or both of the females released eggs in the The series of specimens at night. South of Buenaventura. 1881. Apparently they had recently transformed.29. Choc6: 3984-86. 2935. Specimens Examined COLOMBIA CNHM Antioquia: Puerto Berrio. ECUADOR: 1907. 2920. 30805) there are no pronounced In other specimens (such as blotches on either the back or the tops of the legs.12. we have described here a fresh specimen since the type is old and has faded to a pale in CNHM — brown.118-125. CJG 2249-57.137-146. 4330. 4133. in Narino. JAP 4048-50. . JAP 2798-806. 2996. Condoto. 372. near Lita Station.21. 4058-64. Most of the longpreserved material at hand has faded to a brownish hue.10. collecting bag. JAP 28435.11. p. NARifto: East of Tumaco. east coast of Costa Rica). in fact. 2962. as in some cases. 1915. p. as their head-body lengths range from 19. Pizarro. 30855. USNM 151891. Valle: Buenaventura. Andagoya. 1898. in others. the described specimen. Esmeralda Province: near El Placer. the loreal postocular stripe was bright green. 54772.82. the dark markings are conspicuous on the back and top of the shank but are absent from the thighs. northwest of Lita Station. in March 1962.348 U. MLS 257. west of Santo Domingo de los Colorados. CJG 2249-57 was collected by Goin and on July 13 Medem of Valle. Quibd6.— Brocchi. Two pairs of individuals were clasping. after a torrential rain early in the evening. 2847-8. 4455-61. 37. 1895. Hyla gabbi Cope. 54768-71. Rio Calima above C6r151994.11.— Boulenger. 3044. on the Rio Calima above Cordoba grove. in the state at La Granja. In life dorsal puddles on the the males were calling from little ground color was light brown.S.7 millimeters.

in postaxillary region as wide or a little wider than greatest width of head. disk of fourth toe practically covering the tympanic area. the only other species tympanum of Similisca now known from Colombia. . 34. p. Toes completely webbed. when limbs are laid along the side. when hind leg is adpressed. 22.FROGS OF COLOMBIA 1896.— Liu. Diagnosis. 355. 63893. phaeota. p. broadly U-shaped when viewed from above. separated from each other by an interval equal to about four-fifths their distance from eye. 274 252. when hind legs are bent at right angles to body. 1961. skin of throat and chest smooth. slightly rounded in profile. separated from eye by a distance equal to one-third its own diameter. p. by (tympanum at least two-thirds diameter of eye in phaeota) the smaller the absence of the postocular dark stripe. 1931. 1935. nostrils more lateral than superior. its diameter slightly greater than its distance from nostril. transverse series. palpebral membrane not reticulate. 1960a. third and fifth subequal. —CNHM and by Uraba. Body not elongate. Antioquia. — Goin. moderately projecting. rounded choanae. 1901.— Giinther. p. very broadly cordiform. . 478. p. —COCHRAN —A — Starrett. No patagium. lying close together between the small. p. which is relatively wide and slightly greater than distance between nostrils. that of belly and lower surface of thigh uniformly . Villa Arteaga. its posterior border very slightly free and very shallowly notched. p. a distinct oval inner and a smaller but equally distinct rounded outer metatarsal tubercle. their distance from end of snout equal to about two-fifths their distance from eye. 840. Colombia. Canthus rostralis well defined loreal region concave and somewhat oblique. just reaching to disk of third which snugly covers the tympanic area. interorbital distance slightly less than width of upper eyelid. p. an inner but no outer tarsal ridge. heel reaches slightly beyond tip of snout. Eye moderate but prominent. p. 1960. 1963. Skin of upper parts smooth. heels overlap appreciably. 15. 413. no projecting rudiment of a pollex. 300. Smilisca gabbi. no dermal appendage on heel. p. Nieden. a distinct narrow glandular ridge passing above the tympanum. gabbi may be distinguished from S. 1932. Tympanum very distinct. 1952. 1923. the web reaching the base of disk of all toes.— Taylor.— Gorham. S. p. Vomerine teeth in two short.— Dunn. moderate-sized Smilisca without a dark postocular to only about one-half the diameter and with a tympanum equal of eye. 25.— Werner. the web reaching the middle of penultimate phalanx of third finger fourth finger slightly longer than second. Description. stripe AND GOIN 349 — 351. Snout short. about one-third the diameter of eye. the upper jaw extending a little beyond lower. knee and elbow just fail to meet. Fingers webbed. a low ulnar ridge. 1903. the upper lip flaring out somewhat below it. tongue two-thirds as wide as mouth opening.

350

NATIONAL MUSEUM BULLETIN

U.S.

weak

granular;

no vocal

88

traces of a skinfold across chest; adult gravid female,

Skin of head not co-ossified with

sac.

2

skull, roof of skull

not

exostosed.

Dimensions. Head and body, 52.2 mm.; head length, 17.4 mm.;
head width, 18 mm.; femur, 27.7 mm.; tibia, 29.9 mm.; heel-to-toe,
37.2

mm.; hand,

13.5

Color in alcohol.

mm.

—A

light

brown

dorsally

and cadaverish gray

below. There are small, inconspicuous, scattered, dark

on dorsum and top

of head.

The most conspicuous

brown spots

part of the pattern

a series of distinct light spots along each side between axilla and
Some of the light spots are nearly half the diameter of the
tympanum. These spots are more thickly scattered in region of the

is

groin.

groin. Similar

but

much smaller light spots

surfaces of the thighs.

Very

are present on the posterior

faintly discernible dark crossbands are

on

the dorsal surfaces of the shanks.

We have seen only a single unquestioned specimen of
from Colombia, but we strongly suspect that when adequate
collecting is done around the Gulf of Uraba this species may well be
found to be common.
The two specimens from Sutatenza, in Boyaca, are included as
gabbi only provisionally. Their slender build and small tympanum
separate them from phaeota but a dark postocular stripe and dark,
cruciform blotches on the back make them superficially resemble
that species rather than gabbi. It is possible that they represent an
undescribed race of one of these species but a final decision must be
postponed until fresh material is collected.
Remarks.

S. gabbi

Since we are not acquainted with the frogs in Central America, we
cannot comment on Taylor's assigning of Hyla saluini of Boulenger to
this species.

Specimens Examined

COLOMBIA

CNHM

Antioquia: Villa Arteaga, Uraba,
Boyaca.: Sutatenza, MLS 285, 285a.

COSTA RICA:

Near

Sipurio,

63893.

USNM 30658-9

(cotypes).

Genus Sphaenorhynchus Tschudi
1838.

Sphaenorhynchus Tschudi,
aurantiaca Daudin).

life;

Hyla

lactea T)ax\dm

— Hyla

Moderate to small frogs, bright green or
snout very pointed and projecting in lateral view;

Generic diagnosis.

yellowish in

p. 71 (type species,

male with the external vocal pouch made up of longitudinal folds,
bounded anteriorly and posteriorly by transverse folds; a posteriorly
projecting process on the ischium; reduced number of maxillary teeth
(33 on one side being the greatest number now known in S. aurantiacus
prefrontals not in contact. Very aquatic.
;

.

FROGS OF COLOMBIA

Key
a

1
.

1

a2

ft

.

b2

.

.

to

;

—COCHRAN

AND GOIN

351

Colombian Species of Sphaenorhynchus

Vomerine teeth present and tympanum apparent externally.
A distinct canthal dark stripe from tip of snout to eye
No dark stripe from tip of snout to eye
No vomerine teeth; tympanum not apparent externally
.

.

.

S. aurantiacus

.

.

S. dorisae
.

.

S.

habrus

Sphaenorhynchus habrus (Goin)
Figure 41
1957.

Sphenohyla habra Goin,
1958, p. 121.

p. 18

— Goin, 1961,

1961.

Dryomelictes habra.

1963.

Hyla habra.— Gorham, 1963,

Diagnosis.

—A

(Type

locality, Leticia,

Amazonas, Colombia)

p. 9.

p. 22.

tiny Sphaenorhynchus without an apparent tym-

panum and without vomerine teeth.
The absence of the two above-mentioned

characters serves to sep-

arate S. habrus at a glance from both S. dorisae and S. aurantiacus,

the two other species of the genus

known

to occur in

— UF 8508 (holotype), from

Colombia.

Amazonas, Colombia. Head moderately broad, width about equal to distance from
snout to occiput; snout an obtuse triangle as seen from above, tapering
abruptly to a slightly rounded tip; snout pointed and pronouncedly
projecting as seen from the side; the eyes very dark and thus standing
out prominently against, and through, the milky white skin, but the
Description.

Leticia,

ftk
Figure

41.

Sphaenorhynchus habrus, holotype,
hand, which are X

UF
3)

8508 (X

2,

except foot and

352

U.S.

NATIONAL MUSEUM BULLETIN

2

88

eyes neither particularly large nor protuberant; diameter of eye equal
to distance from eye to tip of snout; nostrils very near tip of snout;
interorbital space equal to about 1% times the diameter of eye; no
external tympanum discernible; upper eyelids, top of head, and dorsum

smooth. The urostyle, vertebrae, and frontoparietal bones bright green
and show clearly through the pale whitish skin of back; the green limb
bones of hind legs plainly seen from ventral surface. No green discernible through skin on front limbs or pectoral girdle. Well-developed
disks on all fingers and toes. Fingers, except first, not particularly
slender; with but the merest rudiment of a web; first finger pronouncedly shorter and more slender than second; fingers 3-4-2-1 in
order of decreasing length.

A

pronounced rudiment

of a pollex.

Toes

moderately slender, all fully webbed except fourth, which is webbed
to penultimate phalanx and has a margin of web to disk; toes 4-3-5-2-1
in order of decreasing length, toes 3 and 5 being nearly subequal.
Heels slightly overlapping when legs are flexed with femora held at
right angles to the body; knee and elbow slightly overlapping when
limbs pressed along the side; heel extending to middle of eye when leg
is pressed along body. No patagium present. Chin, vocal pouch,
venter, and underside of thighs rugose. External vocal pouch a longitudinally folded structure that extends from a horizontal fold at level of
angles of jaws posteriorly to another horizontal fold at level of axillary
region. No well-developed dermal anal flaps. Tongue broadly cordate,
slightly free and slightly notched behind, its width about four-fifths
the width of mouth. No vomerine teeth. Internal nares moderate in

tip

and well separated.
Head-body length, 15.8 mm.; head width, 5.5 mm