WO HARVARD UNIVERSITY Library of the Museum of Comparative Zoology BULLETIN OF THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE, IN CAMBRIDGE VOL. 126 CAMBRIDGE. MASS., USA 1961 - 1962 Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 126 No. 1 SALIENTIA OF VENEZUELA By Juan A. Rivero Institute of Marine Biology and Biology Department University of Puerto Rico Mayaguez, Puerto Rico With One Plate CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM November, 1961 Publications Issued by or in Connection WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE Bulletin (octavo) 1863 — The current volume is Vol. 126. Breviora (octavo) 1952 — No. 145 is current. Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55. Johnsonia (quarto) 1941 — A publication of the Department of Mollusks. Vol. 4, no. 40 is current. Occasional Papers of the Department of Mollusks (octavo) 1945 — Vol. 2, no. 26 is current. Proceedings of the New England Zoological Club (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters "Check List of Birds of the World.'' volumes 1 and 2, 4 and 6 are out of print ; volumes 3, 5, 7 and 9 are sold by the Museum, and future volumes will be published under Museum auspices. Publications of the Boston Society of Natural History The remaining stock of the scientific periodicals of the Boston Society of Natural History has been transferred to the Museum of Comparative Zoology for distribution. Proceedings — Volumes available : 3, 5, 6, 8, 11, 14-17, 20-22, 24-27, 30-34, 37. $4.00 per volume. Occasional Papers: Volume 2, $5.00; Volume 3, $4.00; Volume 4 (1-3), $10.00; Volume 6, $5.00. Memoirs : Requests for some specific memoirs can be filled but no list is available. Bulletin of the Museum of Comparative Zoology AT HAEVAED COLLEGE Vol. 126. No. 1 SALIENTIA OF VENEZUELA By Juan A. Rivero Institute of Marine Biology and Biology Department University of Puerto Eico Mayaguez, Puerto Eico With One Plate CAMBRIDGE, MASS., U.S.A. PRINTED FOE THE MUSEUM November, 1961 No. 1 — Salientia of Venezuela By Juan A. Rivero CONTENTS PAGE Introduction 6 Acknowledgments 6 Historical Eesume 7 Method of Treatment 10 Explanation of Terms and Measurements 11 Key to the Families of Venezuelan Frogs 16 Pipidae 17 Key to Venezuelan Pipa 17 Pipa pipa (Linn6) 17 Pipa parva Euthven and Gaige 18 Buf onidae 19 Key to Venezuelan Bufo 19 Bufo guttatus guttatus Schneider 20 Bufo ceratophrys Boulenger 22 Bufo granulosus granulosus Spix 23 Bufo marinus marinus (Linne) 25 Bufo sternosignatus Giinther 28 Bufo typhonius typhonius (Linne) 29 Bufo typhonius alatus Thominot 31 Leptodactylidae 32 Key to Venezuelan leptodactylid genera 32 Key to Venezuelan Leptodactylus 32 Leptodactylus marmoratus hylaedactylus (Cope) 33 Leptodactylus pentadactylus pentadactylus (Laurenti) 35 Leptodactylus rhodomystax Boulenger 37 Leptodactylus bolivianus Boulenger 38 Leptodactylus mystaceus (Spix) 40 Leptodactylus poeciloohilus dypticus Boulenger 42 Leptodactylus sibilatrix (Wied) 44 Leptodactylus ocellatus (Linne) 45 Leptodactylus podicipinus petersii (Steindachner) 47 Leptodactylus rugosus Noble 50 Lithodytes lineatus (Schneider) 52 Key to Venezuelan Eleutherodactylus 53 Elcutherodactylus cornutus maussi (Boettger) 54 Eleutherodactylus briceni (Boulenger) 56 Eleutherodactylus turumiquirensis sp. n 57 Eleutherodactylus terra ebolivaris sp. n 58 Eleutherodactylus brachypodius sp. n 61 Eleutherodactylus conspicillatus ileamazonicus ssp. n 63 4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Eleutherodactylus stenodiscus Walker and Test 66 Elcuthcrodactylus orocostalis sp. n 68 Eleutherodactylus bicumulus (Peters) 70 Eleutherodact ylus icilliamsi sp. n 72 Eleutherodactylus rozei sp. n 73 Eleutherodactylus urichi (Boettger) 75 Eleutherodactylus reticulatus Walker and Test 76 Eleutherodactylus racenisi sp. n 78 Eleutherodactylus marmoratus (Boulenger) 80 Eleutherodactylus anotis Walker and Test 81 Key to Venezuelan Ceratophrys 83 Ceratophrys calcarata Boulenger 83 Ceratophrys cornuta (Linne) 86 Pseudopaludicola pusilla (Ruthven) 86 (Paludicola fischeri Boulenger) 88 Pleurodema brachyops (Cope) 88 Eupenrphix pustulosus ruthvcni Netting 90 Pseudidae 91 Pseudis paradoxus (Linne) 91 Hylidae 92 Key to Venezuelan hylid genera 92 Key to Venezuelan Eyla 93 Eyla boaus (Linne) 96 Hyla wavrini Parker 97 Eyla albomarginata Spix 98 Eyla granosa Boulenger 99 Eyla geographica geographic-a Spix 101 Eyla crepitans Wied 103 Eyla albopunctata midtifasciata Giinther 105 Eyla loveridgci sp. n 108 Eyla ranireps (Cope) 109 Eyla taurina (Steindachner) 110 Eyla paramica sp. n 112 Eyla jahni sp. n 113 Eyla platydactyla Boulenger 115 Eyla benitezi sp. n 116 Eyla marahuaquensis sp. n 118 Eyla rubra Laurenti 120 Eyla boulengeri (Cope) 121 Eyla baumgardneri sp. n 123 Eyla luteooellata Roux 125 Eyla marmorata marmorata (Laurenti) 126 Eyla tibiatrix tibiatrix Laurenti 127 Eyla tibiatrix ingens (Duellman) 131 Eyla vilsomana meridensis ssp. n 131 Eyla minuta Peters 133 Eyla misera Werner 135 RIVERO : SALIENTIA OF VENEZUELA 5 Eyla orophila planicola Lutz and Lutz 136 Hyla battersbyi sp. n 138 Corythomantis venezolana Mertens 140 Key to Venezuelan Gastrotheca 143 Gastrothcca ovifera (Weinland) 143 Gastrotheca williamsoni Gaige 144 Nototheca pygmaea (Boettger) 145 Key to Venezuelan Phyllomedusa 147 Phyllomedusa bicolor (Boddaert) 147 Phyllomedusa burmeist.eri trinitatis Mertens 148 Phyllomedusa hypocondrialis hypocondrialis (Daudin) 150 Centrolenidae 151 Key to Venezuelan Cockranella 151 Cochranella buclcleyi (Boulenger) 151 Cochranella sp 152 Cochranella fleischmanni (Boettger) 153 Dendrobatidae 153 Key to Venezuelan dendrobatid genera 153 Key to Venezuelan Prostherapis 154 Prostherapis collaris (Boulenger) 154 Prostherapis shrevei sp. n 155 Prostherapis dunni sp. n 157 Prostherapis trinitatis trinitatis (Garman) 158 Prostherapis trinitatis mandelorum (Schmidt) 160 Prostherapis neblina Test 161 Prostherapis alboguttatus (Boulenger) 163 Key to Venezuelan Phyllobates 164 Phyllobates bromelicola Test 164 Phyllobates brunneus (Cope) 166 Dendrobates leucomelas Steindachner 168 Atelopodidae 169 Key to Venezuelan atelopodid genera 169 Key to Venezuelan Atelopus 169 Atelopus oxyrhynchus Boulenger 169 Atelopus cruciger cruciger (Lichtenstein and Martens) 171 Atelopus cruciger vogli Miiller 173 Oreophrynella quelchii quelchii (Boulenger) 174 Oreophrynella quelchii macconnelli Boulenger 175 Banidae 176 Eana palmipes Spix 176 Microhylidae 177 Key to Venezuelan microhylid genera 177 Elachistocleis oralis (Schneider) 177 Otophryne robusta Boulenger 179 Localities 180 References cited 195 b BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY INTRODUCTION The following report contains an account of the frogs of the Venezuelan Republic. It is based mainly on the material collected by the author in Territorio Amazonas and belonging to the Uni- versity of Puerto Rico (U.P.R.) and on specimens deposited in the Museum of Comparative Zoology at Harvard (M.C.Z.), the American Museum of Natural History (A.M.N.H.), the Chicago Natural History Museum (C.N.H.M.), the Museum of Zoology of the University of Michigan (U.M.M.Z.), the United States National Museum (U.S.N.M.) and the Museo de la Universidad Central de Venezuela (U.C.V.), but species reported in the litera- ture and unavailable to the author for examination have also been included. The number of Venezuelan specimens studied comprises a total of about 1250. Twenty-four genera and ninety- six species are represented. The paper as originally written included a description of the physiographical provinces of Venezuela and a short discussion of the fauna that each of them contains. For several reasons, it has been found more convenient to separate the two sections and give priority of publication to the taxonomic portion. It is hoped that the section on zoogeography will follow in the near future. ACKNOWLEDGMENTS I gratefully acknowledge the courtesies extended to me by members of the Venezuelan government, especially Col. Dn. Mi- guel Nucete Paoli, then Governor of Territorio Amazonas, and the secretary Dr. Luis Linares. Mr. Janis Roze, of the Universidad Central de Venezuela, not only has loaned me the frog collection of his institution but has provided valuable information on the localities and habits of the species included. For his courtesy I am more than appreciative. I should be ungrateful if I did not express my appreciation to persons that helped me collect in the field : Mr. Ventura Barnes Jr., leader of the University of Puerto Rico Expedition to Vene- zuela, 1950, whose extensive knowledge of South American tropics made him a valuable advisor and guide ; Dr. Jenaro Maldonado, entomologist and companion of several night searches for frogs and insects ; Dr. Paul Nesbitt, anthropologist of the expedition ; Dr. Hans Baumgartner of Puerto Ayacucho, a great friend of naturalists of Territorio Amazonas and a man with a great desire to help others. From the latter I received a number of specimens as well as his experienced cooperation in the field. RIVERO : SALIENTIA OF VENEZUELA 7 For the loan of specimens used in this study or for information regarding specimens under their custody, I am indebted to Mr. Charles Bogert and Mrs. Bessie Hecht of the American Museum of Natural History, Dr. H. Wermuth of the Berlin Museum, Dr. H. W. Parker, Miss A. G. Grandison and Mr. J. C. Battersby of the British Museum, the late Mr. K. P. Schmidt, Messrs. Clifford Pope and Hyman J. Marx of the Chicago Museum of Natural History, Drs. Norman Hartweg, Charles Walker and James A. Peters1 of the Museum of Zoology of the University of Michigan, Dr. J. Guibe of the Paris Museum, the late Dr. E. R. Dunn of the Philadelphia Academy of Sciences, Dr. Robert Mertens of the Senckenberg Museum and Dr. Doris Cochran of the U.S. National Museum. But my chief obligations for assistance in this work are to the late Dr. E. R. Dunn, Mr. Benjamin Shreve, Mr. Arthur Love- ridge and Dr. Ernest Williams. Dr. Dunn placed at my disposal through letters and through personal conversation his immense knowledge of South American Amphibia, while Mr. Shreve, with his excellent taxonomic eye and accurate memory has helped me solve several problems that I would have been unable to solve myself. Mr. Loveridge and Dr. Williams not only have taken a personal interest in all the phases of my work, but have read the manuscript and suggested some changes. However, for the short- comings of this paper the author assumes full responsibility. Prostherapis dunni, Prostherapis shrevei, Hyla loveridgci and Eleutherodactylus williamsi have been named after these four persons. Hyla benitezi is dedicated to Dr. Jaime Benitez, Chancellor of the University of Puerto Rico, without whose efforts and interest the expedition (during the course of which many of the specimens here reported were collected) would never have been possible. Finally, I am indebted to my wife for the bibliographical work, for typing the manuscript and for her constant help and en- couragement during the course of this work. HISTORICAL RESUME One of the earliest reports of Venezuelan amphibians is that of Lichtenstein and Martens (1856). Recorded as coming from Venezuela are: Notodelphis ovifera ( = Gastrotheca ovifera), i Now at San Fernando Valley State College, Northridge, California. 8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Hyla palmata (-H. boaus), Hyla versicolor (?), Hylodes mar- tinicensis (= Eleutherodactylus gollmeri) ,Crossodaetylus Gaudi- chaudii (?), Cystignathus ocellatus (= Leptodactylus ocellatus), Cystignathus typhonius (= Leptodactylus sibilatrix), Bufo agua (= B. marinus), Bufo Leschenaultii (= B. guttatus) and Bufo strumosus (= B. granulosus). Phrynidium crucigerum (= Ate- lopus c. eruciger) is described from Veragua. In 1863, W. Peters described Hylodes gollmeri on tbe basis of Lichtenstein and Marten's Hylodes martinicensis. I have been unable to determine the origin of this collection, but it appears that many of the specimens were collected by Consul Gollmer in the vicinity of Caracas. Bufo guttatus in all probability came from the southern forests or at least from the Orinoco Delta while Gastrotheca ovi- fera may be the specimen or specimens collected by Appun in Cumbre de Valencia (Ernst, 1877: 281). It was Dr. Dunn's opinion (letters, 1.7.51, 7.10.51) that this collection may have been mixed with one from Panama probably collected by Warschewitz (see remarks on page 173.) Other important collections in the Berlin Museum were made by Appun, Salomon Briceno, Kumurow, Martin, Moritz, Otto, Iiothe, Rosenberg, Wessel and Fisher. Less rich are the collec- tions received from Brandt, Bancard, Eckermann, Effeldt, Ernst, Gundlaeh, Hagenbeck, Hiibner, Klaebisch, Kummer, Mauss, Thivaites, Ursulauf, Valentiner and Liming (vVermuth, letter 25.8.51). A small collection made by Carl Sachs in Ciudad Bolivar and Calabozo was reported on by Peters in 1877. In 1892, 0. Boettger reported on the frogs of the Senckenberg Museum, citing eleven Venezuelan species, all coming from Caracas and collected (except one) by Hiibner and Schlesinger. In a later paper (1893), a group of seven Puerto Cabello (Coll. Mauss) and one "Venezuelan" (Coll. Schultzer) species were added to the list and in 1896, a report was made on 17 species collected by Hiibner in the Alto Orinoco. Most of the Venezuelan specimens reported by Giinther, 1858, and Boulenger, 1882, were acquired by the British Museum from Brandt and Dyson, two Natural History dealers. Dr. Parker informs me (letter, 25.7.51) that he has not been able to discover if Brandt was ever in Venezuela. D. Dyson visited that country in 1851-60 and collected in Colonia Tovar, Valle de Aragua, La Guaira and Cariaco. In 1877, A. Ernst published a book in which he devoted a para- graph to discussing the amphibians of Venezuela, mentioning RIVERO : SALIENTIA OP VENEZUELA 9 four species and two additional genera without citation of species. Stejneger's report on the collections made by Lyons and Robin- son in La Guaira and San Julian came out in the Proceedings of the IT. S. National Museum for 1902. Apart from some separate descriptions and a short report by Boulenger on the frogs col- lected by Brieeno in Merida (1903), little was added to Vene- zuelan amphibiology between 1902 and 1927, when A. Lutz published his "Notas sobre batrachios da Venezuela e da Ilha de Trinidad." Until now this has been the most complete account of the Venezuelan frogs, and although it contains a few errors of interpretation, it is undoubtedly an important contribution and one that has been of great help in the present studies. Dr. Lutz's paper is based mainly on his own collections and observa- tions in the region of Maracay. In the same year Roux published the description of Ilijla luieoccllata on the basis of two male specimens coming from El Mene in the Falcon State (Colls. Kug- ler and Vonderschmitt) . During more recent years other authors have mentioned a few species of frogs. Most important among these are Schmidt (1932), who reported on specimens from Mt. Turumiquire and vicinity (Coll. Blake), Parker (1936), who had specimens from the Upper Orinoco (Coll. Wavrin), Shreve (1947), who reported on the collection made by H. G. Kugler in Falcon State, Aleman (1952 and 1953) who had collections from the region of Baruta, El Ilatillo and Kunana, respectively, and Walker and Test (1955) who described three species of Eleutherodactylus from Rancho Grande (Coll. Test). The most noteworthy and until now unreported collections in the American museums are those of Baker (U.M.M.Z. ; Carabobo, Yaracuy, Tachira); Barnes (C.N.H.M. ; Yaracuy, Aragua, D. F.) ; Beebe (U.S.N.M. ; Aragua, Monagas) ; Briceho (Me- rida); Carriker (U.S.N.M.; U.M.M.Z., Merida); Cherrie (U.S.N.M.; Bolivar) ; Holt (U.S.N.M.; Aragua, Miranda, D. F., Terr. Amaz.) ; Mondolfi and Vivas Berthier (U.S.N.M.; Aragua, Miranda, D. F., Falcon, Guarico) ; Osgood and Osgood and Con- over (C.N.H.M.; Merida) ; Pinkus (U.M.M.Z. ; Roraima Region) ; Rosenberg (Merida-); Tate and Tate and Carter (A.M.N.II. ; Turumiquire, Duida and Roraima regions) ; Vogl (U.S.N.M., C.N.H.M.; Aragua); Schultz (U.S.N.M.; Maracaibo Basin); Weber (M.C.Z. ; Bolivar) ; and Williamson, E. and J. (U.M.M.Z. ; Carabobo, Yaracuy, Tachira). 2 Represented in most museums. 10 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY In 1942, E. Kohl published his "Fauna Descriptiva de Vene- zuela," which was revised in 1949. A few species of frogs are included and although there are several errors, the hook fur- nishes common names and contains two records otherwise re- ported only by Ernst. The Universidad Central de Venezuela has a good representa- tion of native frogs, which has been mostly collected by the director of the museum, Dr. Racenis, and Mr. Janis Roze, a young instructor and enthusiastic curator of Herpetology. Most of the specimens come from Aragua, Distrito Federal and Guarico. Except for the types and paratypes, the tag numbers of the U.C.V. specimens should be considered provisional. METHOD OF TREATMENT The Venezuelan frog fauna, as known today, comprises 24 genera and 93 species. The discussion of each of them includes the Venezuelan synonymy and a description that, except when otherwise stated, is based strictly on Venezuelan material. Where specimens were not available, the original descriptions have been used. The contents of these have been slightly modified to suit my purpose but in no case (and no matter how poor the origi- nal) has the description been altered in meaning or the original remarks modernized in any way. The group of specimens used in the writing of the description in each case is listed following the synonymy. Usually the speci- mens chosen for the description are from a single region. This has been done to insure the homogeneity of the sample described. Following the description of the species, measurements (in mm.) of a male and female are given, not necessarily of the largest, but of the best preserved specimens. Measurements of most of the specimens studied were recorded but it has not been considered desirable to publish them in the present report. The list of localities includes those for the material deposited in the museums mentioned in the introduction of this paper as well as those given in the literature. When a specimen was not examined to my complete satisfaction (because it was tempo- rarily missing from the shelves or for some other reason) its number is followed by an asterisk. The heading "Additional Localities" is used in each case to include records of specimens that have not been used in the description. Following the list of localities is the range of the species. This includes the range within Venezuela and also other countries RIVERO : SALIENTIA OF VENEZUELA 11 where the forms under consideration are said to occur. This last is based on the literature and on the specimens deposited in the different museums. Although representatives of many of the species occurring' in the surrounding countries were carefully examined, in some instances it has been necessary to accept the records taken from the files. A list of most of the localities mentioned in this paper, and a description of those where frogs have been collected is given at the end of the taxonomic section. EXPLANATION OF TERMS AND MEASUREMENTS The following is a list of the terms and measurements used in the text. A short explanation of amphibian terminology is given to help students who are beginning in this field, while Spanish equivalents are supplied in the belief that they may be of some value to the trained English-speaking specialist as well as to the Latin American student having little knowledge of English. 1. Head (cabeza). The head may be as long as broad (tan larga como ancha), longer than broad (mas larga que ancha) or broader than long (mas ancha que larga). The length is measured between the tip of the snout and the posterior mar- gin of the tympanum, the breadth at the greatest distance between the tympana. When the tympanum is hidden the measurements are taken from or between the angles of the mouth. As the ratio of length to breadth can be easily found from the measurements given for each species, the character is only mentioned in the description when the difference from other species in this regard should be emphasized. In some cases, as in the genus Leptodactylus in general, and espe- cially in L. bolivianus and L. occllatus, the ratio of length to breadth is quite variable although the proportion given is the usual one. 2. Crests or ridges (crestas o aristas). Ridges that occur on the heads of some bufonids and hylids. The subnasal ridge (arista subnasal) is usually the semicircular prolongation of the canthal crest around the nostrils (Bufo granulosus). The canthal crest (cresta cantal) runs along the canthus rostralis; the orbitotympanic or supratympanic (orbitotim- panica o supratimpanica) lies above the tympanum, and the orbital (orbital) around the orbit. The latter may be divided into pre-, supra-, post- and infraorbital. 12 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY M\ o / 0 \ Oy ^ 0--0 m n °sP O 0\/^ 1 o o O Q> s / n ^ 7\ W A 7\ Fig. 1. a-g, shapes of snout; h-k, shapes of tongue; 1-s, vomerine teeth; t-z, canthus and loreal region. See text pp. 11-16. 3. Snout (hocieo). The snout may be semicircular (semicircu- lar) as in Figure la, e.g. Centrolene buckleyi; almost semi- circular (casi semicircular) as in Figure lb, e.g. Hyla mar- morata; rounded (redondeado) as in Figure lc, e.g. Hyla RIVERO : SALIENTIA OF VENEZUELA 13 granosa or Leptodactylus rugosus; subovoid (subovoide) as in Figure Id, e.g. Leptodactylus pentadactylus ; subelliptical (subeliptico) as in Figure le, e.g. Leptodactylus bolivianus; or acuminate (acuminado, aguzado) as in Figure If, e.g. Leptodactylus sibilatrix. It may project over or overhang the mouth (saliente) in a shark-like fashion as in Figure lg, e.g. Leptodactylus sibilatrix; be truncate (truncado), e.g. Prosthcrapis trinitatis, or swollen and elevated in front (hinchado y elevado al frente), e.g. Hyla granosa and boans. 4. Derm of the head involved in cranial ossification (piel de la cabeza comprendida en la osificacion craniana). Easily recog- nized by determining if the skin is free from the cranium. Gastrotheca ovifera is a good example. 5. Tongue (lengua). With respect to form, the tongue may be circular or rounded (circular o redondeada) as in Figure Ih, e.g. Centrolene buckleyi; cordiform (cordiforme) as in Figure li, e.g. Hyla tibiatri.r; pyriform or spatulate (piri- forme o espatulada) as in Figure lj, e.g. Bufo guttatus; or oval (ovalada) as in Figure Ik, e.g. Leptodactylus bolivianus. With respect to its breadth, it may be broad (aneha), e.g. Hyla lover idgei, or narrow and with parallel edges (estrecha y con margenes paralelos), e.g. Bufo granulosus. It is free (libre) only when attached to the anterior part of the mouth. and adherent (adherente) when completely attached to the floor. When indented (indentada) behind, it is said to be emarginate (emarginada), nicked or notched (escotada, in- cisa), and when lacking the indentation, entire (entera). 6. Vomerine teeth or odontoids (dientes u odontoides vomeri- anos). In a series of two on the palate, in the vicinity of the posterior nares. They may be distinct (distintos) as in most species, indistinct (indistintos), e.g. Pleurodema brachyops, or absent as in Bufo, Prostherapis and some hylids. They may form different designs as in Figures 11 to lp ; between (entre) the choanae (choanas) as in Figure lq ; behind (detras) them as in Figure lr; or behind and between (detras y entre) them as in Figure Is. 7. Canthus rostralis (canto rostral). The ridge or angle formed between the eye and tip of the snout by the outer edges of the snout bones. It may be angular (angular) as in Figure It, e.g. Bufo guttatus (a right angle is implied) ; obtusely angular (angular obtuso) as in Figure lu, e.g. Hyla boans; rounded (redondeado) as in Figure lv, e.g. Hyla minuta, 14 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY indistinct (indistinto), or absent (ausente) where no external angle is formed and the snout would present a smooth, semi- circular appearance if seen in cross-section. 8. Loreal region (region loreal, region frenal, mejilla [Es- pada]). The area bounded anteriorly by the nostrils, pos- teriorly by the eyes, dorsally by the canthus and ventrally by the upper lip. It may be straight or vertical (recto o verti- cal) as in Figure lw, e.g. Bufo guttatus; almost vertical (casi vertical) as in Figure lx, e.g. Bufo typhonius; little sloping or oblique (poco inclinado u oblicuo) as in Figure ly, e.g. Hyla misera and crepitans; and oblique or sloping (obli- cuo o inclinado) as in Figure Iz, e.g. Hyla granosa. It may also exhibit a depression (depresion) in which case it is concave or excavated (concavo o excavado) or be completely flat or non-concave (piano o no concavo). On occasions it may be slightly convex (un poco convexo). 9. Interorbital space equal, shorter or broader than an upper eyelid (espacio interorbital igual, mas eorto o mas ancho que un parpado superior). The interorbital space is measured in the middle, between the two upper eyelids, the eyelid by depressing it slightly and measuring from the origin to the outer margin. When the space is said to be equal to the eyelid it can be expected to be slightly narrower or broader in some specimens. As definite numbers used in the compari- son (e.g., interorbital space 1% times broader than an upper eyelid) are of no value in practice, and may even obscure the importance of other characters, they are used as little as possible in the text. 10. Eye diameter equal, shorter or longer than distance between eye and nostril (diametro del ojo igual, mas corto o mas largo que la distancia entre el ojo y la nariz). The diameter of the eye is measured between the two corners, the distance between eye and nostril, between the anterior corner of the eye and the posterior of the nostril. When the diameter of the eye is as long as the snout (tan largo como el hocico) it is implied, of course, that it is also longer than the distance between eye and nostril. Only in genera where the character is variable and important, the comparative distance between nostril and eye and nostril and tip of the snout is given, e.g., nostril closer to the eye than to the tip of the snout (nariz mas cerca del ojo que de la punta del hocico). RIVERO : SALIENTIA OP VENEZUELA 15 11. Metacarpal and metatarsal tubercles (tuberculos metacar- picos y metatarsicos [Espada]). Since they are present in most species, they are only mentioned in the description where they are larger or smaller than usual or when they are modified in such a way as to be useful in the determination of the species. In some cases one or both of the metacarpal or metatarsal tubercles are absent. 12. Snbarticnlar tubercles (tuberculos subarticulares, nndillos o pelotillas infra-articulares [Espada]). Found under the articulations of the digits and toes. Their mention is avoided if they lack any distinctive feature. 13. Web (palmeadura [Espada], membrana interdigital, mem- brana natatoria). Loveridge's system of measuring the amount of webbing by giving the number of free phalanges has been adopted here as it has the advantage of being standard, a feature that is absent in the other system (i/o webbed, etc.). Since different authors use different fingers or toes or all of them to determine the amount of webbing, it is difficult to make a general comparison, but in general it can be said that Boulenger's % webbed fingers have li/o free phalanges on the outer finger, his %, 2 and his %, 23/2. When present, the disk contains the outer phalanx. For measuring the number of free phalanges, the fingers and toes are placed close together and the extension of the web is determined by considering the middle, not the mar- gins of the membrane. The web of the first finger is that between the first and second fingers, of the second, that be- tween second and third, of the third, that between third and fourth and of the fourth, that between third and fourth also but in relation to the fourth. 14. Lateral fringes of toes and fingers (margen cutanea de los dedos de los pies y las manos). This character seems to be quite valuable in some cases. When said to be faintly indi- cated, the fringes may be completely absent in some speci- mens. On the other hand, a species that does not normally show lateral fringes may show some indication of them in a badly desiccated state. Allowance should be made for such cases. In a few species, e.g., Leptodactylus podicipinus, there are lateral flaps that may go around the toe for 14 of its cir- cumference. 15. Tarsal fold (repliegue tarsico). A cutaneous fold or ridge that extends usually obliquely along part or all the length of 16 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY the tarsus. In some species, e.g., Pseudopaludicola pusilla, the tarsal fold begins with a tubercle in the middle of the tarsal segment. 16. Metatarsal fold (repliegue metatarsieo). A slight skin fold or close line of tubercles tbat extends along the outer margin of the metatarsal segment and outer toe. 17. Hidden portions of the hind limbs (porciones ocultas de las extremidades posteriores) . Those parts of a frog's hind limb that cannot be seen from above or below when the animal is in the resting position. 18. Heel of the adpressed hind limb extends to the shoulder, the tympanum, the eye, the nostril, the tip of the snout (la ex- tremidad posterior extendida a lo largo, el tobillo llega a el hombro, el timpano, el ojo, la nariz, la punta del hocico). Obtained by stretching the bind limb anteriorly and deter- mining the place to which the tibio-tarsal articulation ex- tends. Sometimes a valuable character although it should not be interpreted too closely. A broken limb bone alters the heel extension completely. 19. Secondary sexual characters (caracteres sexuales secunda- rios), as the hooks or rugosities on the inner finger of some males, the breast spines of Leptodactylus pentadactylus, the chin spines of Leptodactylus rugosus, the swollen arms of Leptodactylus ocellatus, the external vocal pouches of many species, the longer arms of the male Atelopus oxyrhynchus, the snout ridge of Leptodactylus mystaceus, the tubercular dorsum of the male Hyla teturina, etc. Most of these charac- ters are only shown by the males during the breeding season. Key to flu Families of Venezuelan Frogs I. Vertebrae convex anteriorly, concave posteriorly (opisthocoelous) ; sacrum fused with the urostyle. Eustachian tubes opening through one orifice into the pharynx; tongue and eyelids absent PIPIDAE II. Vertebrae concave anteriorly, convex posteriorly (procoelous) ; sacrum articulated with the urostyle by two condyles. A. Right and left half of the pectoral girdle overlapping, movable (arciferal). 1. No small cartilage (intercalary) between the last and penulti- mate phalanges; terminal phalanges simple or T-shaped; trans- verse processes of the sacral vertebrae (sacral diapophysis) cylindrical or dilated ; belly smooth or granular. a. Omosternum absent, parotid gland present, upper jaw not toothed BUFONIDAE RIVERO: SALIENTIA OF VENEZUELA 17 b. Omosternum present, parotid gland absent or small and inconspicuous; upper jaw toothed LEPTODACTYLIDAE 2. A small piece of cartilage (intercalary) between the last and penultimate phalanges; transverse processes (sacral diapophy- sis) of the sacral vertebrae generally dilated; belly granular; toes generally webbed. a. Terminal phalanges T-shaped, astragalus and calcaneum fused CENTEOLENIDAE b. Terminal phalanges claw-shaped, astragalus and calcaneum not fused HYLIDAE 3. An accessory phalanx in digits; terminal phalanges simple (Pseudis), or claw-shaped (Lysapsus) ; transverse processes (sacral diapophysis) of sacral vertebrae cylindrical; toes webbed PSE TJDIDAE B. Eight and left half of the pectoral girdle fused in the middle, immovable (flrmisternal). 1. Omosternum present; sternum cartilaginous; fingers and toes with dermal scutes on the disks DENDBOBATLDAE 2. Omosternum absent; sternum bony; fingers and toes without dermal scutes on the tips ATELOPODIDAE III. First seven vertebrae procoelous; 8th biconcave, sacral, biconvex (di- plasciocoelous), articulated with the urostyle by two condyles. A. Teeth present, palate normal EANIDAE B. Teeth absent, palate with two dermal ridges in front of the pharynx MICEOHYLIDAE PIPIDAE Key to the Species of Pipa Recorded from Venezuela I. Two lateral flaps at the angle of the mouth ; one large metatarsal tubercle pipa II. Lateral flaps at the angle of the mouth absent; no metatarsal tubercles parva Pipa pipa (Linne) Rana pipa Linne, 1758, Syst. Nat., ed. 10: 210: Surinam. Pipa dorsigera Ernst, 1877, Flora y Fauna de Ven. : 281. Pipa pipa Dunn, 1948, Amer. Mus. Novit., no. 1384: 9. Pipa americana Eohl, 1949, Fauna Descr. de Ven., ed. 2: 395, fig. 177. No Venezuelan material examined. 1 (M.C.Z. 6127) Trinidad. 3 (M.C.Z. 1244) Dutch Guiana. Description. Head flattened, openly triangular but with a blunt tip ; nostrils transversely elongated, sometimes with a 18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY slightly elevated piece of skin at the outer edges ; a short tentacle under the tip of the snout; mouth very large, with two distinct lateral flaps at its angles ; one or more short, pointed appendages above the small eyes and sometimes another pair on each side of the snout under the upper lip ; fingers long and slender, each ending in four tips ; metacarpal tubercles absent ; thighs very short ; a very prominent inner metatarsal tubercle but no outer ; toes broadly webbed to the tips. Skin above, shagreened and with small round tubercles and lateral line organs. Below, rugose. Measurements. Snout-vent S 135, 9 120. Additional Localities. A group of petroleum geologists (Lopez, Davey y Rubio, 1946 : 123) have observed this species in the Delta Region. Ernst and Rohl do not give any special local- ity. Range. Eastern Venezuela. Trinidad and the Guianas to Matto Grosso; Colombia to Peru and probably Bolivia. Remarks. According to Rohl the common name in Venezuela is "sapo de celdas." Pipa parva Ruthven and Gaige Pipa parva Euthven and Gaige, 1923, Occ. Papers Mus. Zool. Univ. Mich., No. 136 : 1 : Sabana de Mendoza, Venezuela ; Noble, 1925, Amer. Mus. Novit., No. 164: 2; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 41, pi. viii, fig. 1; Dunn, 1948, Amer. Mus. Novit., No. 1384: 8. Protopipa parva Carvalho, 1939, Bol. Biol., 4: 397, figs. 6b, 6c. 1 (U.M.M.Z. 55743, type) Sabana de Mendoza. 4 (U.M.M.Z. 57444-57447, prtps.) Sabana de Mendoza. 1 (M.C.Z. 9012, prtp.) Sabana de Mendoza. 47 (U.S.N.M. 115770-816) 20 km. w. Rosario, iii.42. Description. Head depressed ; snout rather pointed and pro- jecting beyond the mouth ; nostrils transversely elongated, ending laterally in two projecting tips; eyes small, their diameter much shorter than distance between eye and nostril ; interorbital space broad ; no dermal appendages at the tip of the snout nor in any other place on the head ; arms relatively short ; fingers short, with a small basal web, each ending in four short appendages; meta- tarsal and subarticular tubercles absent ; toes fully webbed, their tips pointed ; tarso-metatarsal articulation of the adpressed hind limb extends to the eye or between eye and nostril. Skin above, uniformly covered with keeled tubercles. Below, tubercular ex- cept on the lower lip. RIVERO: SALIENTIA OF VENEZUELA 19 Color. Grayish brown, tinged with yellowish and with obscure darker markings. Below, brownish yellow. Measurements. Snout-vent $ 29, 9 34 ; head breadth $ 13, 9 9.5 ; head length S 7, 9 6 ; femur 6 12.5, 9 12,3 ; tibia & 13.5, 9 13. Habits. Females with swollen backs were found to be gravid. Apparently some hormone ( ?) stimulates swelling prior to spawn- ing. Some specimens only 11 mm. in length had swollen backs. Additional Localities. El Mene, in a roadside puddle that drains into Rio Cocuiza (U.S.N.M. 115818-20) ; Lagunillas (U.S.N.M. 11757-8) ; Rio San Juan (U.S.N.M. 115767-9) ; Zulia (U.S.N.M. 128845, juv.). Range. The arid and semiarid Maracaibo Basin and the Fal- con Region. Northeastern Colombia. Remarks. This species represents the only frog that is ap- parently endemic to the arid and semiarid region of northwestern Venezuela. It seems to be most closely related to Pipa aspera of the Guianas, but apparently there is complete discontinuity of their ranges. It is possible that in previous times an ancestral form occupied all the area from the Maracaibo Basin to the Guianas and that the range was broken by the changes that oc- curred during the Ice Age. Living in a medium of high specific heat, this mostly aquatic animal was able to persist in the Mara- caibo Basin while failing to survive in the Coastal Range owing to the peculiar conditions existing there at the time. I agree with Dunn (1948) in including the genera Pipa, Proto- pipa and Hemipipa in the single genus Pipa. BUFONIDAE Key to the Species of Bufo Reported from Venezuela I. Head without bony ridges.1 A. Snout truncate; canthus angular; a wedge-shaped, skin-covered ridge in front of the eye g. guttatus II. Head with bony ridges.1 A. Upper eyelid with a hornlike dermal appendage. 1. A lateral fringe of pointed tubercles; snout with a triangular tip c.eratophrys B. Upper eyelid normal. 1. A subnasal ridge; tympanum in contact with the descending ramus of the postorbital ridge ; nostrils directed upward ; heel never extending to parotid g. granulosus 2. No subnasal ridge; tympanum not usually in contact with the descending ramus of the postorbital ridge; nostrils lateral or almost so; heel generally extending to parotid or beyond. i This character does not usually work for young specimens. Such juveniles should be closely examined for the presence or absence of the supraorbital ridge. The use of the other key characters may also be necessary. 20 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY a. Angles of the jaw not projecting. 1. Tarsal fold strong; snout not prominent, gradually in- clined downward toward the front m. marinus 2. Tarsal fold absent; snout prominent, not gradually in- clined downward toward the front. a. Palms and soles spinulous ; metatarsal fold absent sternosignatus b. Palms and soles not spinulous ; metatarsal fold present typhonins aJatus b. Angles of the jaw strongly projecting; palms and soles not spinulous; metatarsal fold present t. typhonins Bufo guttatus guttatus Schneider Bufo guttatus Schneider, 1799, Hist. Amphib., 1: 218: "India Orientali"; Boettger, 1896, Ber. Senckenb. Naturf. Ges.: LIV. Bufo Leschenaultii Lichtenstein and Martens, 1856, Nomencl. Rept. Amphib. Mus. Berol. : 42. 7 (U.P.R. 30-36) Pto. Ayacucho, v.50. 4 (U.P.R. 37-40) Tapara, vi.50. 5 (U.P.R. 42-46) Raudal de Dios, v.50. 2 (U.P.R. 41, 47) La Culebra, 1,000 ft., v.50. 1 (M.C.Z. 17708) Sn. Fernando, 1895. Description. Snout truncate, as long as, or very slightly longer than, the eye diameter; tongue pyriform, entire or slightly nicked behind ; eye diameter greater than distance between eye and nostril; interorbital space flat, about twice as broad as an upper eyelid ; canthus angular ; loreal vertical, flat ; tympanum distinct, y2 the eye diameter ; a wedge-shaped, skin-covered ridge in front of the eye ; other head ridges absent ; sometimes the slight but thick supratympanic eminence appears to be an anterior exten- sion of the parotid ; parotids large, distinct, pitted, extending on the sides as far down as the lower margin of the tympanum; first finger much longer than second; fingers and toes swollen at the tips, with a thick but not very distinct lateral fringe; subarticular tubercles large, single, rounded, that of the first finger larger than the others ; tarsal fold fairly prominent ; meta- tarsal fold absent; inner metatarsal tubercle very prominent; toes taken in order from first to fifth exhibit the following phalanges free of web: 1*4, 1, 2, 3%, 2; heel of the adpressed hind limb extends to the posterior margin of the parotid. Skiu on the posterior part of the dorsum with numerous flat and very RIVERO: SALIENTIA OF VENEZUELA 21 distinctive warts; flanks and limbs warty; a series of 3 or 4 transverse rugae at the nape ; a loose fold of skin on each side of the body from parotid to groin ; ventral surfaces of the belly and thighs rugose and with flat warts or granules. Color. Above, olive or wood-brown with orange brown warts ; limbs, flanks and sides of the head blackish or brownish gray ; parotids particolored. Below, brownish gray, uniform or more usually with scattered, round, white spots; lower jaw generally margined with white dots ; limbs occasionally exhibiting dark blotches on the ventral surfaces. Measurements. Snout-vent $ 47, ? 85; head breadth $ .1.5, 5 24.5 ; head length $ 14.1, $ 23 ; femur $ 16, $ 27 ; tibi'i $ 14, 9 26. Habits. Like B. typhonius with which it is usually found, this species apparently prefers the leaf -covered floors of wooded areas. Unlike B. t. typhonius, however, it seems to be more par- tial to lower grounds, the species ceasing to occur at about 400 m. on the Marahuaca region, where B. typhonius is still abundant. The six largest adult females (U.P.R. 30-36) were collected at night while they were crossing a road after a heavy rain. Their ovaries were completely undeveloped. Additional Localities. Upper Orinoco (Boettger, 1896) ; Vene- zuela (Licht. and Mart., 1856). Range. The Venezuelan Guayana and probably the Delta Re- gion. The Guianas to northern Brasil and with all probability southeastern Colombia. Remarks. The label with the type specimen (Berlin 3517) in- dicates that it was collected in Surinam by Bloch. Although the locality records are not sufficient to draw any final conclusions, I consider Bufo glaberrimus and Bufo guttatus sufficiently close structurally to be regarded as conspecific. To avoid the use of a racial name until the intergradation is confirmed appears as un- desirable as to make premature use of the trinomial. These two forms are apparently not separated by any physical or faunal barrier, their ranges are allopatric and the differences they ex- hibit appear to be of racial value. Bufo guttatus guttatus is apparently very sensitive to changes of latitude and altitude (see "Habits" and "Range" above) and it is possible that these factors have contributed in differentiating glaberrimus although this animal is found as low as 500 m. in Colombia and Ecuador. It is distinguished from typical guttatus 22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY by the absence of preorbital ridges and in having a longer hind limb, the heel usually extending to the anterior half of the paro- tid. The skin is smooth in both forms up to a size of 35 or 40 mm. when the flat, rounded warts appear. These are generally more orange in guttatus than in glaberrimus. I have not discovered any difference in webbing between the two forms. The rose spot in the inguinal region described by Giinther for B. glaberrimus is found in a specimen from Macanal, Colombia (M.C.Z. 15059, 68 mm.) but not in those from Sarayacu, Ecuador (M.C.Z. 19603-12 plus 56 dupl., largest 53% mm.). The latter are generally dark in the anterior part of the belly and throat but the white round spots that occur in some B. guttatus are some- times present in B. glaberrimus. M.C.Z. specimens 3807-8, 2968 (largest 5iy2 mm.) from Sali- dero, N. W. Ecuador undoubtedly represent a race, distinguished by the variegated dorsum, greater amount of web and densely dotted ventral surface. It is not impossible that they represent the immature condition of the recently described Bufo blombergi Myers and Funkhouser, which might also be included in the same Rassenkreis. Boulenger (1882) mentions a Bufo guttatus gut- tatus of 177 mm. (Demerara), so it appears that the species in general may attain considerable size. Besides its normally large size, however, Bufo blombergi is distinguished from glaberrimus by several minor differences, and geographically it is disjunct, being separated by the whole Andean Range. I believe Bufo anderssoni Melin (Taracua, Uaupes R., Brasil) is a synonym of Bufo guttatus guttatus. This would appear from the ridge in front of the eye shown in one of the figures. Bufo ceratophrys Boulenger Bufo ceratophrys Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 319, pi. xii, fig. 2 : Ecuador. 1 (U.P.R. 253, juv.) Temiche, 4050 ft., v.50. Description. Snout with a pointed triangular tip ; nostrils somewhat raised laterally ; tongue narrow, entire ; eye diameter greater than distance between eye and nostril ; interorbital space broader than an upper eyelid ; canthus well defined ; loreal al- most straight ; tympanum hidden ; probably slight supraorbital and supratympanic ridges ; parotids moderate, subovate, super- ficial ; first finger a little shorter than second ; a moderate meta- tarsal fringe to the tip of the outer toe ; subarticular tubercles large ; toes webbed at the base only ; heel of the adpressed hind RIVERO: SALIENTIA OF VENEZUELA 23 limb extends to the eye. Skin above covered with small tubercles ; three distinct tubercles at the angle of the mouth and several others at the ventral base of the arm; upper eyelid with a tri- angular dermal appendage ; a lateral fringe of tubercles from parotid to groin. Color. Above, brown with minute dark points and a dark spot on one of the sides of the dorsum ; fringes along the sides of the body black-edged; limbs crossbarred. Below, uniformly light brown. Measurements. Juv., snout-vent. 12 mm. Habits. The only specimen was collected at night on the leaf- covered floor of very humid forest. Range. Only known from Marahuaca in Venezuela. Eastern face of the Andes of Ecuador. Remarks. Although very small, the specimen shows sufficient characters for its proper allocation to the species. It differs from two Ecuadorean specimens examined (M.C.Z. 19601-2, Canelos to Maranon) and from the original description in having the first finger shorter than the second, a hidden tympanum and no well-developed lateral papillae. The last two characters are prob- ably a sign of immaturity as the smaller of the Ecuadorean specimens is similar in these respects. So far as I am aware, the species has not been reported from the Amazonian forests between the Guayanan Cerros and the Andes. It represents the first Guayanan frog showing Andean affinities and a confirmation of similar findings in plants, birds and mammals. More abundant and adequate material of Bufo ceratophrys may possibly show some racial differentiation from the Andean form. Bufo granulosus granulosus Spix Bufo granulosus Spix, 1824, Spec. Nov. Testud. Ran.: 51, pi. xxi, fig. 2: Bahia ; Boettger, 1892, Kat. Batr. Samni. Mus. Senckenb. : 39 ; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 42, pi. viii, figs. 5, 6; Parker, 1936, Bull. Mus. Roy. Hist. Nat, Belgique, 12: 1. Bufo strumosus Lichtenstein and Martens, 1856, Nomencl. Rept. Amphib. Mus. Berol. : 42. Bufo granulosus granulosus Miiller and Hellnuch, 1936, Wissen. Ergeb. Deutsch Gran Chaco Exped. : 13. 13 (U.P.R. 16-28) Sn. Fernando de Atabapo, vi.50. 1 (U.P.R. 29) Pto. Ayacucho, vi.50. 24 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Description. Snout prominent, slightly longer than the eye diameter ; nostrils directed upward ; tongue narrow, entire ; eye diameter greater than distance between eye and nostril ; inter- orbital space about IV2 times broader than an upper eyelid ; canthus curved, denned by a horny ridge that extends anteriorly as a semicircle around the nostril ; loreal slightly oblique ; tym- panum moderate, the anterior border in contact with the post- orbital ridge, the posterior margin usually obscured by granules ; orbits encircled by a low but distinct, horny ridge ; a horny labial and a short supratympanic ridge ; parotids of moderate size but not distinctly marked ; first finger not extending beyond second ; a rounded palmar and a smaller and less distinct inner meta- carpal tubercle ; subarticular tubercles single, the second of the third finger double ; edges of fingers and toes serrated ; tarsal and metatarsal folds absent; toes taken in order from first to fifth exhibit the following phalanges free of web: 1%, 1%, 2,-2y2, 3%, iy^-2; heel of the adpressed hind limb does not reach the parotid. Skin above, tubercular, spiny on the sides and limbs. Male with a subgular vocal sac, and dark rugosities on the inner side of the first digit. Color. Above, light brown or yellowish gray, with irregular dark gray markings ; head ridges black. Below, yellowish gray, usually marbled with darker gray on the breast and anterior part of the belly. Measurements. Snout-vent $ 46.7, 9 52 ; head breadth 8 14.5, 9 16 ; head length $ 12.1, 9 14 ; femur $ 10, 9 13 ; tibia $ 13.5, 9 17. Habits. Most of the specimens were caught in amplexus in the shallow waters of a rain pool in the savannas of San Fer- nando. It is strange that while the B. marinus collected in the same locality were heavily infested with Amblyomma dissimile, not one of the B. granulosus was attacked by this tick. Of three stomachs examined, one was empty, one contained five small snails, mud, two small stones and a piece of wood. The other had one ant, one small hemipteran and three or four pellets of unidentifiable remains. Additional Localities. Arabopo (U.M.M.Z. 85131-2, 85133 [2], 85134[2]); Caracas (U.C.V. 100; Boettger, 1892); Espino (U.C.V. 68); Lake Maracaibo (C.N.H.M. 3016[2]); Maracay (U.S.N.M. 97193-5; Lutz, 1927); Parmana (U.C.V. 10); Pto. Cabello (Licht. and Mart., 1856) ; Sanariapo (U.S.N.M. 80641- 50) ; Upper Orinoco (Parker, 1936) ; Venezuela (Licht. and Mart., 1856). RIVERO : SALIENTIA OF VENEZUELA 25 Range. The arid and semiarid Maracaibo Basin, the northern coastal belt, the arid and semiarid sections of the Coastal Range, the Llanos and the northern half of the Venezuelan Guayana. Colombia and the Guianas to northern Argentina. Remarks. The specimens from Arabopo (4000 ft.) have very prominent crests and perhaps should deserve racial recognition. Bufo marinus marinus (Linne) Rana marina Linne, 1758, Syst. Nat., ed. 10: 211: America (Bestricted to Surinam by Miiller and Hellmich, 1936). Bufo agua Lichtenstein and Martens, 1856, Nomencl. Kept. Amphib. Mus. Berol.: 42; Giinther, 1858, Cat. Batr. Sal. Brit. Mus.: 65. Bufo marinus Peters, 1877, Monatsb. Akad. Wissensch. Berlin: 460; Bou- lenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 315; Boettger, 1892, Kat. Batr. Samm. Mus. Senckenb.: 39; 1893, Ber. Senckenb. Naturf. Ges.: 40; 1896, Ber. Senckenb. Naturf. Ges.: LIV; Stejneger, 1902, Proc. U. S. Nat. Mus., 24 : 180 ; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 42; Aleman, 1952, Mem. Soc. Cienc. La Salle, U\ 27, fig. 3. Bufo marinus marinus Schmidt, K. P., 1932, Zool. Ser. Field Mus. Nat. Hist., 18: 159. 7 (U.P.R. 48-51, 76-7, 89) Sn. Fernando de Atabapo, vi.50. 1 (U.P.R. 52) Chorro Chupadero, iii.50. Description. Snout slightly longer than the eye diameter; tongue oval or pyriform; eye diameter greater than distance between eye and nostril; interorbital space about iy2 times as broad as an upper eyelid; canthus well defined, slightly curved and sloping toward the front ; loreal short, almost vertical ; tym- panum not very distinct on the posterior margin, y2 to % the eye diameter ; head ridges well defined, the infraorbital extending obliquely to the angle of the jaw, the parietal variably distinct ; a few tubercles behind the angle of the jaw; parotids large, pitted, obliquely subtriangular ; a longitudinal series of tubercles from hand to elbow ; first finger longer than second ; a very large, flat palmar and a smaller inner metacarpal tubercle ; subarticular tubercles generally single or incompletely fused, the second of the third finger double ; tarsal fold strong ; metatarsal absent ; toes taken in order from first to fifth exhibit the following phalanges free of web : 1, 1 to 1%, 2, 3 to S%, V/2 to 1% • the web is ex- tended to the tips as distinct lateral fringes ; heel of the adpressed hind limb extends to the parotid. Skin above and on the flanks covered with warts of irregular size ; upper eyelids and sides of the head tubercular ; ventral surfaces granular, the granules be- ing sometimes tipped with a dark corneus matter. Male with a rugosity on the inner side of the first digit. 26 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Color. Above, brown with irregular lighter and darker mark- ings. Below, light brownish or yellowish, generally marbled with gray. Measurements. Snout-vent $ 44, 9 79 ; head breadth $ 15, 9 26 ; head length $ 14, 9 24 ; femur $ 16, 9 32 ; tibia $ 17, 9 35. Habits. B. marinus seems to be fairly common along the Ori- noco River where its voice was often heard at night. Its presence was not detected in the Cunucunuma nor in the Upper Duida and Marahuaca regions. Several of the specimens collected in Sn. Fdo. de Atabapo were infested with the tick Amblyomma dissimile. One of the animals had been so emaciated by the infestation that for a while it was taken for some other form. Male and female specimens were collected in a shallow rain pool together with B. granulosus. While most of the B. granu- losus were in amplexus, the ovaries of the adult B. marinus were completely undeveloped. Additional Localities. Rio Albarregas (A.M.N.H. 10501) Ara- bopo (U.M.M.Z. 85120[3]) ; Barrancas (M.C.Z. 19916, juv.) ; Ba- ruta (Aleman, 1952) ; Calabozo (Peters, 1877) ; Campo del Lago, Lagunillas (U.S.N.M. 115692-8) ; Caracas (Boettger, 1892) ; Ciu- dad Bolivar (M.C.Z. 19114, 19 juv.) ; Cocollar, Cumana, Cumana- coa (Schmidt, 1932) ; Duida Region (A.M.N.H. 32962) ; El Periquito (U.C.V. 89) ; El Valle (U.S.N.M. 128850-2) ; Espino (U.C.V. 50) ; Kunana, Perija Mts. (Aleman, 1953) ; La Fria (U.M.M.Z. 55585*); La Guaira (U.S.N.M. 22537-8, 27800-1, 27803-6, 27796); Macuto (U.C.V. 77-8*); Maracay (U.M.M.Z. 52707-8*, Lutz, 1927) ; 30 km. of Maturin (U.C.V. 76) ; Petare (U.S.N.M. 121170) ; Pto. La Cruz (U.S.N.M. 121171-2) ; Pie del Cerro U.S.N.M. 121168) ; Puerto Cabello (Boettger, 1893) ; Sn. Antonio (U.S.N.M. 836119) ; Sn. Carlos (U.S.N.M. 83610) ; Sn. Fernando de Atabapo (U.S.N.M. 80651); Sn. Julian (U.S.N.M. 27807); Sta. Catalina (M.C.Z. 19915, 4 juv.); Sta. Elena, Bolivar (U.M.M.Z. 85121); Sta. Lucia (U.S.N.M. 121169); Tanaguarena (U.C.V. 36); Turgua (Aleman, 1952); Turgua (U.S.N.M. 129265) ; Upper Orinoco (Boettger, 1896) ; Venezuela (C.N.II.M. 43664? [3-6 juv.]; U.M.M.Z. 55583-4*; Lichtenstein and Martens, 1856 ; Giinther, 1858 ; Boulenger, 1882). RIVERO : SALIENTIA OF VENEZUELA 27 Range. Probably all the physiographical provinces of Vene- zuela including the Andes to an elevation of 1,600 m. Texas to northern Brasil. Remarks. No constant difference in size of parotid between specimens from northern and southern Venezuela was observed. While those of the southern animals are generally of a large size, some of the northern specimens (U.S.N.M. 121165, 121171, 27800) have glands that rival in volume any of those of the Amazonian group, with the possible exception of the larger southern speci- men, an animal from San Antonio having a snout-vent length of 142 mm. The coloration of U.S.N.M. 83610 (131 mm.) is unusual: the ventral surfaces, the under sides of the parotids, the lower jaw and the posterior part of the thighs are bright yellow and dark marbled while the throat is profusely infuscated, almost black. Dr. Cochran and I have agreed that this specimen represents B. marinus but in coloration it is not very different from a British Museum Pto. Cabello specimen that has been referred to B. crucifer (Blgr., 1882: 317, a or b). I have examined this latter animal but have excluded the species from the fauna of Vene- zuela until its presence is confirmed by further material. The male Bufo marinus is more tubercular above, the tubercles are more spiny, and the color is usually more uniform than in the female. Cochran, 1955, treats Bufo paracnemis and Bufo ictericus as distinct species, on the ground that the two overlap in part of their ranges (Rio de Janeiro and Sao Paulo). On the east, B. paracnemis is said to occupy the intermediate area between north- ern B. marinus and southern B. ictericus, occurring in Brasil in Minas Gerais, Bahia and Pernambuco. To the west the southward extension of Bufo marinus is un- certain, as records may refer to any of the related forms. How- ever, it does not appear impossible that Bufo marinus may have given rise to paracnemis on the east and ictericus on the west, and that these two forms may have come together after their inde- pendent origin from a common ancestor. Anyway, there does not appear so far to be sufficient evidence against considering paracnemis a race of Bufo marinus and on that basis the tri- nomial is used here. Bufo marinus andinensis Melin, 1941, was described from Roque, Peru. Bufo pythecodactylus Werner is considered a synonym of Bufo marinus marinus. 28 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Bufo sternosignatus Gunther Bufo sternosignatus Gimther, 1858, Cat. Batr. Sal. Brit. Mus. : 68, pi. vi, fig. c. : Pto. Cabello, Venezuela; Cordova, Mexico (restricted to Vene- zuela by Boulenger, 1882); Boulenger (part?), 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 323; Boettger, 1892, Kat. Batr. Samm. Mus. Senckenb. : 39; Gunther, 1901, Biol. Centr. Amer., Bept. and Batr.: 247; Nieden, 1923, Das Tierreieh Anura. I: 182; Lutz, A., 1927, Mem. Inst. Osw. Cruz, SO: 38, 42, pi. viii, figs. 3, 4. Bufo typhonius sternosignatus Shreve, 1947, Bull. Mus. Comp. Zool., 99 : 535. 2 (M.C.Z. 17812-3) Distrito Acosta, 29. 1 (M.C.Z. 26151) Cerro Cosine, x.39. 3 (M.C.Z. 26152, 25976-7) Pauji, v. 45. 3 (U.M.M.Z. 55690-3) Sn. Esteban. 1 (U.M.M.Z. 55694) Kio Bejuma. 2 (U.M.M.Z. 55695-6) Rio La Mona. 1 (U.M.M.Z. 55697) Bet. Valencia and Caserio Silva. 1 (U.C.V. 16) Carayaca, xii.51. 1 (U.C.V. 28) Cerro Avila, viii.51. Description. This species is very similar to Bufo typhonius alatus, from which it can be distinguished by the following characters: snout not very prominent; orbital ridges slight, the supratympanic usually well developed, the parietal absent or very indistinct ; parotids subovate, not generally pointed pos- teriorly ; hands and feet spinulous; the subarticular tubercles not different from the other spinules of palms and soles ; metatarsal Hold absent ; heel of the adpressed hind limb extends to the posterior half of the parotid. Skin above and below very granu- lar, occasionally coarse and spinulous especially on the sides of the body and venter. A lateral fringe of tubercles is found in some but not all individuals. Color. This animal is usually of a reddish color when alive. In alcohol it turns brown and many specimens present a colora- tion very similar to that of B. typhonius. Some specimens are peculiarly marked with yellow and brown below. Measurements. Snout-vent $ 41, 9 49; head breadth $ 14, 9 16; head length $ 12, 9 14; femur $ 16.5, 9 18; tibia $ 16, 9 18. Habits. Apparently a forest animal. Additional Localities. Caracas (Boettger, 1892) ; El Limon (U.S.N.M. 121173-4) ; Mamo, La Guaira (Lutz, 1927) ; Puerto Cabello (Gunther, 1858; Boulenger, 1882) ; Venezuela (Gunther, 1858; Boulenger, 1882; Nieden, 1923). RIVERO : SALIENTIA OF VENEZUELA 29 Range. The Coastal Range and Falcon Region. Reported from Colombia by Peracca, 1914 (Cafetal Camelia, nr. Angelopolis, 1820 m.), Werner, 1916 (Canon de Tolima), and Boulenger, 1882 (Bogota). These records probably refer to B. typhonius alatus. Dunn (1944) does not mention this species in his "Herpe- tology of the Bogota Area" but he includes Bufo typhonius in the actual fauna of Bogota. Remarks. This species is very close to Bufo typhonius alatus with which it has been often confused and with which it perhaps interbreeds (though not freely) in part or all of its range. Gen- erally it is well characterized (U.M.M.Z.) but specimens are found (M.C.Z. and U.C.V.) that are very similar to B. typhonius, especially in coloration and general appearance. In all these cases the palm and fingers are spinulous (not a sexual character) and the metatarsal fringe absent. The supraorbital ridge is also more prominent in B. typhonius and presents a pleated appearance in the region where a parietal ridge would join the supraorbital. The upper eyelid thus appears to be lower on the sides in B. typhonius than in B. sternosignatus. If a perpendicular line were to be drawn from the upper edge of the supraorbital ridge to the outside, the upper eyelid would be mostly above the line in sternosignatus and below it in typhonius. Bufo typhonius typhonius (Linne) Fana typhonia Linne, 1758, Syst. Nat., ed. 10: 211 : America. Bufo typhonius Boettger, 1896, Ber. Senckenb. Naturf. Ges. : LIV. Bufo typhonius typhonius Leavitt, 1933, Copeia : 8. 7 (U.P.R. 4-10) Slopes Mt. Marahuaca, 4000 ft., v.50. 7 (U.P.R. 1-3, 11-12, 14-15) Upper Cunucunuma R., v-vii.50. 1 (U.P.R. 13) Anaben, Colombia, vi.50. 1 (U.S.N.M. 83949) Yapacana, iv.21. Description. Snout prominent, with a more or less distinct vertical ridge at the tip, from the level of the nostrils to the edge of the jaw ; tongue narrow, entire ; eye diameter slightly greater than distance between eye and nostril; interorbital space twice as broad or broader than an upper eyelid ; canthus well defined, carved ; loreal almost vertical and a little concave ; tympanum moderate, oval, % the eye diameter; preorbital and descending ramus of the postorbital ridge slight or absent ; supraorbital ridge strong, anteriorly merging with the canthal, posteriorly with the very prominent supratympanic crest ; parietal ridge distinct, forming a basin with the parotid ; parotids prominent but small 30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY and superficial; edge of the upper jaw studded with minute acu- minate tubercles ; two knob-like protuberances at the angle of the jaws; first finger not extending beyond second; tips of the fingers and toes swollen, their sides serrated ; tarsal fold absent ; subarticular tubercles large, usually single or fused ; a row of closely set, small spinules along the outer edge of the metatarsal segment and fifth toe ; toes partially webbed, the web reaching at least to the penultimate phalanx in all the toes except the third, where it does not extend beyond the proximal phalanx; heel of the adpressed hind limb extends to between parotid and tym- panum. Skin above and on the sides of the head tubercular; tubercles spiny on the flanks and limbs ; a fringe of pointed tubercles from anterior border of parotid to groin ; ventral sur- faces granular, some of the granules spiny on the pectoral region. Color. Above, brown, uniform or with irregular lighter and darker markings ; from tip of snout to end of urostyle a yellowish vertebral stripe of varying width, on either side of which, espe- cially in the sacral region, roundish black spots may be present or absent ; flanks below the lateral fringe, occasionally lighter or darker than the dorsum. Below, yellowish brown, plain, infuscate or spotted; limbs with broad, more or less distinct crossbars. Measurements. Snout-vent $ 42, 9 61 ; head breadth $ 14, 9 21 ; head length S 14, 9 18; femur