BREVIOIA Museum of Comnparatlve Zoology Cambridge, Mass. 31 December, 1968 Number 306 A NEW SPECIES OF ELEUTHERODACTYLUS (AMPHIBIA, SALIENTIA) FROM THE GUAYANA REGION, EDO. BOLIVAR, VENEZUELA Juan A. Rivero Abstract. A new species of Eleutherodactylus, E. pulvinatus, is de- scribed from the wet region of La Escalera, Estado Bolivar, Venezuela. The species is characterized by its small but distinct tympanum, poorly defined or absent vomerine teeth, large, transversely oval disks which are larger than the tympanum, a W-shaped marking on the occiput and non- spiny nuptial pads. The possible relationship between this frog and the genera Eupsophus, Syrrhophus, Niceforonia, Trachyphrinus and Phryno- pus is discussed and evaluated. Among the specimens recently collected by the author in the region of La Escalera, Estado Bolivar, Venezuela, there are seven specimens of a small and peculiar leptodactylid whose generic allo- cation has been somewhat difficult and problematical. In this species, the vomerine odontoids are very indistinct; in some indi- viduals they appear to be absent altogether. As in Eleutherodac- tylus, the disks are well developed and the terminal phalanges are T-shaped, but there are two well-developed glandular pads on the inner side of the first digit, a character which does not seem to have been described in that genus. It has been customary to allocate all Eleutherodactylus-like. forms without vomerine teeth to the genus Syrrhophus (type local- ity, San Antonio, Texas) and all those with vomerine teeth but without disks to Eupsophus (type locality, Chile). In 1932, Parker violated convention by describing Eupsophus wettsteini (type local- ity, Pongo, Peru, 2000 m) as "the first species that has been de- scribed as belonging to the genus Eupsophus in which vomerine teeth are lacking ,, (p. 344). Even a superficial examination reveals that Andean species of Syrrhophus are not closely related to the animals that are referred 2 BREVIORA No. 306 to that genus in Central America and the United States. Whether or not they have vomerine teeth, most of the Andean animals seem to be closely related and should perhaps be called Eupsophus until a more thorough study can be undertaken and their relationships established. It is evident that the three groups of Eupsophus (Thoropa, wettsteini-peruanus, and the Chilean forms) mentioned by Cei ( I960. 1962) constitute different evolutionary lines, one of which {wettsteini-peruanus) appears to continue along the Andean Cordillera to Venezuela, where Eupsophus ginesi is known from glacial streams in the paramos oi the Merida Andes. It is now USUall) recognized that Thoropa is a separate genus, and perhaps the wettsteini-peruanus-ginesi group should be recognized as an- other, in which case. Oreobates Jimenez de la Espada, 1872. might appear to be the appropriate name. Yet. although Oreobates was described from Quijos (whence the name quixensis), and this name has been interpreted to mean Quito (Peters. 1955: 347; Gorham 1966: 117). Espada himself states: "It occurs in the Oriental province, formerly Quijos. in Ecuador, at the foot of the Cordillera" ( \ crtcbrados del \ iajc del Pacifico, 1875: 104). Specimens from the Napo-Pastaza region in the M.C.Z. correspond to the description o\' quixensis, which looks more like a Thoropa than a Eupsophus. Two new genera, each based on a single individual from the high elevations of the Colombian Andes, were described by Coin and Cochran in 1963. The first. Nicejoronia, has distinct vomerine teeth, no disks and no external tympanic disk; the other. Trachy- phrinus, has no vomerine teeth but has small disks (and T- shaped phalanges) and a well-defined tympanum. In general physiognomy, both of these genera resemble the group of frogs here considered, and Trachyphrinus does not look at all like Syr- rhophus, the genus in which it would be included if its classifica- tion were to be based on conventional morphological characters. But all kinds of combinations of characters have been described in the high elevation species from Peru to Venezuela. Thus, in 1873. Peters described Phrynopus peruanus (now called Eupso- phus, fide Gorham. 1966) with vomerine teeth and tympanum, but apparently without disks. The same combination is found in Borborocoetes columbianus Werner (now Eupsophus, fide Gor- ham. 1966). Paludicola festae Peracca (now Syrrhophus, fide Gorham. 1966). and Eupsophus ginesi Rivero. However, in Pal- udicola simonsii Blgr. (now Syrrhophus, fide Gorham, 1966). Syr- rhopus montium Shreve and Syrrhopus juninensis Shreve (both now Syrrhophus, fide Gorham. 1966) there are no vomerine teeth. 1968 NEW SPECIES OF ELEUTHERODACTYLUS 3 external tympanic disk, or disks. In Eleutherodactylus whym- peri (Blgr.) and E. surdus (Blgr.), on the other hand, vomerine teeth and disks are present but the tympanum is not apparent externally, while in Eupsophus wettsteini there are no vomerine teeth or disks but there is a small tympanic disk. Tra- chyphrinus myersi and Syrrhophus areolatus Blgr. are alike in having disks and external tympana (small and ill defined in S. areolatus) but no vomerine teeth. It is not the purpose of this paper to enter into the many prob- lems pertaining to this interesting group (or groups?) of frogs. But these digressions have been necessary because of the diffi- culty of allocating the new species described in this paper. If this group of Andean frogs is found to constitute a natural assemblage, perhaps the name Phrynopus can be used for it. It is also possible that it may constitute two or more evolutionary lines, in which case the genera suggested by Goin and Cochran may be found useful. It appears to me, however, that this group, or most of it, constitutes a latitudinal extension of Chilean and Argen- tinian Eupsophus along belts of similar climatic conditions in the Andes. If, as has been reported, some of the species are limited to paramos in the Andean Cordillera, their present diversification is to be expected. In a group where the tympanum-vomerine odon- toids-disks can occur in so many combinations, separation into various genera is hardly convenient and may actually be very confusing. A number of "Syrrhophus" have been described from the low- lands or at most, from elevations below 800 m in Ecuador. Such is the case, for example, with S. chalceus (Peters), S. calcaratus Andersson, and S. coeruleus Andersson. S. chalceus has also been reported by Lutz and Kloss, 1952, from Iuarte, along the Rio Uaupes. As the largest of the species described was 30 mm (5. coeruleus), all the others ranging from 18 to 29 mm, it is not im- probable that the lack of vomerine teeth can be attributed to immaturity in some of them. It is possible, however, that they represent another line of Eleutherodactylus without vomerine teeth (since in this genus disappearance of the vomerine teeth does not seem to be a difficult process) or that they are in some way con- nected with the Eupsophus group from Chile and/or the more northern Andes. Eleutherodactylus pulvinatus, the species described here, does not have the rotund physiognomy of the Eupsophus group, and its distribution does not suggest relationship. Nuptial pads have been described for some members of the Andean group (Parker, 1932: 4 BR! MORA No. 306 363), hut most of the species do not seem to have them. In the Chilean species (Cei, l l >o2). on the other hand, and in Thoropa (Cochran. 1955: 91), thumb and sometimes breast and axillar spines have been described for those species in which males arc known. Similar modifications have been described for Crossodac- tylodes Cochran (type localit) Macae. Rio de Janeiro), a inono- typic genus with hidden tympanum and the vomerine teeth represented onh b\ a roughened ridge. In view o\' the foregoing considerations, it has been found preferable to describe the new leptodactylid in the genus Eleutherodactylus, pending a more complete studv o\ its anatoim and relationships. I I I l llll RODA< H I. US Pill VINATUS sp. n. Type. M.C.Z. 64741, ■' , Paso del Danto, Region de la Esca- lera around 1400 m above San [sidro, road from II Dorado to Sta. Elena de Uairen. Estado Bolivar, Venezuela. Coll. J. A. Riveroand J. Pulido, :<> March 1968. Diagnosis. A small Eleutherodactylus with small but distinct tympanum, vomerine teeth poorl) defined or absent; two non-spin) nuptial pads on the inner side o\ the first digit o\ males; large trans- versel) oval disks, larger than the tympanum; first linger shorter than second; adpressed heel extending to between eye and nostril: and a W -shaped marking behind the occiput. Description. Mead as long as broad; snout short, subtriangular; tongue free and slightl) nicked behind: vomerine odontoids faintl) indicated as two irregular elevations well behind ami between the small choanae; eye diameter slightk greater than distance between eye and nostril but shorter than snout: interorbital space as broad as an upper eyelid; canthus rostralis rather indistinct. curved; loreal region onl) slightK sloping, concave; tympanum dis- tinct. ' .* the eve diameter; a Hat. indistinct supratympanic fold; a line of small, whitish tubercles from elbow to wrist; two distinct. oval, metacarpal tubercles; a glandular pad at the base o\' the inner side of the first finger and another one distal and slightK dorsad to this; first finger shorter than second, its disk much broader than the phalanx; all fingers tree: larger disks larger than the tympanum; a small. Hat. whitish tubercle at the distal portion o\ the tibial segment: a distinct and prominent inner and an indis- tinct outer metatarsal tubercle: toes free, with prominent subarticu- lar tubercles: heel of the adpressed hind limb extending to between eye and nostril. Skin above, shagreened. more coarsely tubercular in the tympanic area and anterior flanks. Below, finely granular on 1968 NEW SPECIES OF ELEUTHERODACTYLUS 5 throat and anterior part of the belly; more distinctly granular on posterior part of the belly and thighs. A very large subgular vocal sac with a triangular anterior end; a space (about 2 mm) between tip of the jaw and anterior edge of sac. A "frenulum" in the an- terior tip of the sac continues anteriorly almost to the tip of the jaw. Color. Above, brownish gray, with a dark brown interorbital bar, a black supratympanic streak, a W-shaped marking on the anterior part of the back, and some large blotches and small spots on the posterior two-thirds of the back; on the flanks the blotches continue ventrally, forming oblique, wavy bars that are separated by light gray areas; anterior and posterior part of the thighs uniform brown, this color extending as bars across the gray color of the upper part of the thighs; limbs and fingers with cross-bars and blotches; upper lip with triangular blotches on a gray background; no canthal streak. Ventral surfaces dirty white, darker on the throat and distal portion of the limbs. Measurements (in mm). Snout-vent 26.1; head breadth 9.8 (between tympana); head length 10; femur 13; tibia 14.1; foot 1 1. Paratypes. There are six paratypes (U.P.R.-M. 1 02214-02219), all coming from the type locality. All are males, and the size varies from 26.1 mm to 23 mm. In some of the smaller specimens the canthus seems to be less defined than in larger specimens, and a cross-section of the snout would be more or less rounded if it were not for the slight concavities of the loreal areas. The heel of the adpressed hind limb extends to the nostril in only one speci- men, but the skin of the dorsal surfaces is more granular than in the type, at least in two examples. The belly and thighs are uni- formly granular, the throat more finely so in most specimens. The vocal sac does not form the anteriorly triangular pouch that it forms in the type, but it is possible that it has been dis- placed, as a similar condition can be noticed in several specimens. The W-shaped marking is present in all specimens, but the oblique bands of the flanks are less distinct in some, although a trace of them can be noticed in all. Vomerine odontoids cannot always be seen, but they can be felt with a needle. They seem to be irregu- lar in both size and position, and in some specimens seem to be almost totally absent. Habits and relationships. All the specimens were collected at night, calling from low bushes ( 1 to 3 ft. from the ground) along the rocky bank of the road at an elevation of approximately 1400 m. At dusk, their calls seemed to come from under the rocks 1 University of Puerto Rico — Mayagiiez. 6 BREVIORA No. 306 lining the almost vertical road bank, but none could be collected at the tune. Later at night, their calls were traced and recorded and the various specimens collected. In the dark the Erogs are yellow or cream, but they turn dark gray in the daytime. Their voices were not heard much below the elevation where all the specimens were obtained. No other species of Eleutherodactylus was col- lected or heard in this area. Only scattered low bushes, some soil bromeliads and. occasion- ally, thickets of lilaceous plants occur in the rather open areas where E. pulvinatm was collected. There were permanent and temporary streams everywhere, as the region is probablv one ol the rainiest in Venezuela. 1 The forest. 50 to 60 meters from the road, is a rather open montane forest, usuallv with man) epiphytic bromeliads. orchids, and araeeans. Most of the soil is a white quartziferous sand, which is covered in the forest by a layer of dead leaves and other organic matter. Although mam Eleutherodactylus have W-shaped markings on the back, the distinct markings ol E. pulvinatUS remind one oi the condition o{ E. W-nigrum (Boettger), which, however, is a species from more than 3000 m in the Amies o\ Ecuador. Other species. from relativel) adjacent regions, are: /•.'. inguinale Parker (New River, Guyana), E. beebei Cochran (Kartabo), E. pliciferus (Blgr.) and E. ramagii ( Blgr. ) (Iguarasu, Pernambuco, Brasil), / crepitans Bockermann (San Vicente, Matto Grosso), E, car- valhoi B. Lutz (Itacoai R.. NW Brasil). E. peruvianus (Melin) (Roque, Peru). E. vilarsi (Melin) (Taracua, Rio Uaupes), / rosmelinus Gorham (Rio Uaupes). /.. melini Bokermann (Roque, Peru). /:". altamazonicus Barbour and Dunn (upper Amazon) and /■.. marmoratus (Blgr.) (Mt. Roraima). In all these species there are well-defined vomerine teeth, and nuptial pads have not been described for am. E. inguinale has on obtusely angular canthus, a broad interorbital space, much broader than the upper eyelid, and a black bordered, inguinal ocellus; E. beebei Cochran is a large species 85.5 mm in length; in E. pliciferus and E. ramagii the first linger is longer than the second; E. crepitans, from drv forest, is similar to /.. conspicillatus, and its tympanum is % the eve diameter; E. carvalhoi does not have a tympanum; E. peru- vianus and E. vilarsi are 50 and 39 to 41 mm. respectively, and may be synonyms of E. conspicillatus; E. rosmelinus has a depressed head, disks VS the size of the tympanum and first finger 1 According to a report in a local newspaper, engineers working on the road from El Dorado to Sta. Elena could only work 60 to 80 days of the year, and even those were not always sunny days uithout any rainfall. 1968 NEW SPECIES OF ELEUTHERODACTYLUS 7 longer than second; E. melini has an indistinct tympanum and small disks; E. altamazonicus has a depressed and oval snout and a barely distinguishable tympanum; and E. marmoratus has a rounded snout as long as the orbital diameter, dark-edged marb- lings on the head, and a small size (only topotypical specimen: 19 mm). 1 From the lowland "Syrrhophus" mentioned before, E. pulvinatus can apparently be distinguished by the nuptial pads it has in the inner side of the first finger. Other differences are as follows: In S. chalceus (Peters), from Pastaza River, the first finger is longer than the second; in S. coeruleus Andersson, from eastern Ecuador, the canthus is distinct and there are two pairs of dorsolateral folds; and in S. calcaratus Andersson,- from near Archidona, 800 m, there is a canthal streak, and the disk of the first finger is only slightly broader than the phalanx behind it. Acknowledgments. The author wishes to acknowledge all the courtesies and attentions extended to him by the Instituto Vene- zolano de Investigaciones Cientificas of Caracas, and most espe- cially by Dr. Gilberto Rodriguez, Head of the Hydrobiology Department, who went out of his way to foster the investigations resulting in this and other papers. During the course of this study the author was employed by and received all kinds of assistance from the I VIC. For the trip to La Escalera, the author relied on an invaluable assistant, Mr. Juan Pulido, always faithful and always willing to help during the difficult situations that often arise during such trips. Major Luis Alfonso Godoy, Commander, Engineering Corps at sector A of the road from El Dorado to Sta. Elena, was extremely helpful in providing facilities and assistance. As always, the author is greatly indebted to Dr. Ernest Williams, not only for permitting the use of the facilities of the Museum of Comparative Zoology at Harvard, but also for reading the manu- script critically and for discussing various taxonomic and evolu- tionary problems. Mr. Benjamin Shreve has also read the manu- script and suggested changes. The author, however, assumes full responsibility for the shortcomings of this paper. 1 Frogs collected in Mt. Marahuaca by the author (Rivero, 1961: 80) and pronounced conspecific with E. marmoratus by Parker, do not repre- sent E. pulvinatus. However, further comparisons should be made when more specimens of E. marmoratus become available. - Among the group of frogs here discussed, the species name calcaratus was first used by Giinther, 1881, for Cacotus calcaratus, now considered a synonym of Eupsophus roseus. 8 bki vior \ No. 306 REFERENCES VNDl RSSON, 1 \KS (i \HKII 1 1945. Batrachians from Easl Ecuador collected 1937. 1938, b) Wm. Clarke-Macintyre and Kolt" Blomberg. Arkiv Zool. 37<2): 1-88. B.XRUOl R. I IIOM \s. \ND I R. l)l NN 1921. Herpetological novelties Proc Biol Soc. Wash. 34: 157-162. linkl KM \NN. \\ I RM R 1958. A preoccupied name of a neotropical frog, genus Eleuthero- dactylus. Herpetologica I 1. pt. 2: 95. 1965. Tres novos batraquios da regiao central de Matto Grosso, Brasil (Amphibia. Salientia). Rev Brasil. Biol. 25(3): 257-264 BOI l I NG1 R, Cil I iRGl \l HI Rl 1882. Catalogue ol the Batrachia Salientia in the Collection of the British Museum, ed. 2. I ondon, Kvi -♦ 503 pp. 1888. 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