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list of those reported from the Virgin Islands. Ann. New York Acad. Sci., 10:1-160.

Schwartz, A. 1958. Four new frogs of the genus Eleutherodactijlus ( Lepto- dactylidae) from Cuba. Amer. Mus. Novitates, (1873): 1-20.

Schwartz, A. 1964. Three new species of frogs ( Leptodactylidae, Eleuthero- dactijlus) from Hispaniola. Breviora, (208): 1-15.

Schwartz, A. 1965. Variation and natural history of Eleuthewdactylus rtithae on Hispaniola. Bull. Mus. Comp. Zool., 132:479-508.

Schwartz, A. 1966. The relationships of four small Hispaniolan Eleuthewdac- tylus (Leptodactylidae). Ibid., 133:369-399.

Schwartz, A. 1967. Frogs of the genus Eleutherodactijlus in the Lesser An- tilles. Stud. Fauna Curacao and other Carib. Isl., 24 (91): 1-62.

Schwartz, A. 1969. The Antillean Eleutherodactijlus of the auriculatus group. Ibid., 30(114 ):99-115.

Schwartz, A. 1973. Six new species of Eleutherodactijlus ( Anura, Leptodac- tylidae) from Hispaniola. J. Herp., 7:249-273.

Schwartz, A. 1976. Two new species of Hispaniolan Eleutherodactijlus ( Lep- todactylidae). Herpetologica. 32:163-171.

Schwartz, A., Thomas, R. 1975. A checklist of West Indian amphibians and reptiles. Carnegie Mus. Nat. Hist., Spec. Publ., (1):1-216.

Shreve, B., Williams, E. E. 1963. The frogs, pp. 302-342. In Williams, E. E., Shreve, B. and Humphrey, P. S., The heipetology of the Port-au- Prince region and Cona\e Island, Haiti. Parts l-II. Bull. Mus. Comp. Zool., 129:291-342.

Smith, H. M., Taylor, E. H. 1948. An annotated checklist and key to the Amphibia of Mexico. Bull. U. S. Natl. Mus., (194): 1-1 18.

Stejneger, L. 1904. The herpetology of Puero Rico. Ann. Rept. U. S. Natl. Mus., 1902:549-724.

Stuart, L. C. 1963. A checklist of the herpetofauna of Guatemala. Misc. Publ. Mus. Zool. Univ. Michigan, (122): 1-150.

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OCCASIONAL PAPERS

of the

MUSEUM OF NATURAL HISTORY

The University of Kansas ^^y Lawrence, Kansas ..^ 4 ic~7

NUMBER 62, PAGES 1-10 DECEMBER 21, 1976

DESCRIPTION OF A NEW SPECIES OF

TELMATOBIINE FROG, TELMATOBIUS

(AMPHIBIA: LEPTODACTYLIDAE),

FROM THE ANDES OF NORTHERN CHILE

By

Alberto Veloso M/ and Linda Trueb"

Five species of anurans are known to occur in the Andes of northern Chile between latitudes of 18° and 23° S (Capurro, 1950, 1953, 1955; Noble, 1938; Sclimidt, 1954); these are: Bufo spinulosus (Wiegmann), Pleurodema marmorata (Dumeril and Bibron), Tel- matohius marmoratus (Dumeril and Bibron), T. peruvianus (Wieg- mann) and T. halli (Noble). As a result of extensive field work in the Department of Arica in northern Chile, it is now possible to add a previously undescribed species to the anuran fauna of this area. This stream-inhabiting frog herewith is assigned to the genus Tehnatobius. Its discoveiy contributes to our understanding of ecological and systematic relationships among the anurans of northern Chile, especially those that occur at high altitudes in the central cordillera of the Andes.

The first author wishes to express special gratitude to The Uni- versity of Kansas which served as his host institution, and William E. Duellman of the Museum of Natural History, who graciously provided facilities for his research. This research was made pos- sible through funds provided by a General Research Grant (No.

^ Associate in Herpetology, Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045 and Associate Professor, Departamento de Biologia Celular y Genetica, Universidad de Chile, Casilla No. 6556, Santiago, Chile.

" Adjunct Curator, Di\'ision of Herpetology, Museum of Natural History and Adjunct Assistant Professor, Department of Systematics and Ecology-, The Uni- versity of Kansas, Lawrence, Kansas 66045, U.S.A.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

3272 to W. E. Duellman) from the Graduate School of The Uni- versity of Kansas, and travel to the United States was funded by the Programa Multinacional de Genetica, OEA, Chile. We are indebted especially to our colleague Jaime Pefaur for his continued interest and unremitting encouragement, and herewith dedicate this new species to him with appreciation.

Telmatobius pefauri, new species Figures 1 and 2

Hohtype. — The University of Kansas Museum of Natural His- tory (KU) 159836, an adult female from Murmuntani, 3200 m, Departamento de Arica, Provincia de Tarapaca, Chile (latitude 18° 2V S, longitude 69° 27' W), collected on 23 November 1972 by Alberto Veloso M.

Diagnosis. — Telmatohius pefauri is a moderate-sized species that can be distinguished from its congeners by the following combina- tion of characters: 1) hind limbs long-tibio-tarsal articulation of adpressed limb lying anterior to eye; 2) snout rounded in dorsal profile, depressed in lateral profile; 3) mandibular arch not prognath- ous; 4) nostrils barely protuberant; 5) tympanic annulus concealed externally and incompletely developed under the skin; 6) maxillaiy and premaxillary dentition embedded in mucosal labial folds; 7) dentigerous processes of prevomers small, horizontally oriented, bearing a moderate medial separation and located at the mid-level between large choanae; 8) eyes dorsolateral with a distinct anterior orientation; 9) tarsal fold well developed; 10) maxillaiy dentition present; and 11) prevomerine dention present.

Description. — Adult female, moderately large and with robust body, 75 mm in snout-vent length ( Fig. 1 ) ; males not known. Head large, its length 37 percent of snout-vent length, depressed and broader than long (head length only 82% of head width); in dorsal profile, snout broadly rounded and in lateral profile, long and sloping from the orbital region to a subacuminate temiinus; lips thick and glandular externally and with maxillaiy and premaxillary teeth embedded in mucosal labial folds in the buccal cavity; nostrils small, scarcely protuberant, located approximately equidistant from the tip of the snout and the ventral margin of the lip and closer to the orbit than to the tip of the snout; internarial region flat; loreal region concave; canthus rostralis straight and poorly de- fined; tympanum absent externally but remnant of tympanic an- nulus present beneath skin. Supratympanic fold poorly developed and diminutive in size and length, extending from posterior border of orbit to approximately mid-point between end of mouth and axilla, where it is obscured within lateral skin folds of body. Eye large (diameter approximately 29% of head length), positioned far

A NEW SPECIES OF TELMATOBIUS FROM CHILE

Fig. 1.— Holotype of Tclmatobius pefauri, adult female (KU 159836). Line equals 1 cm.

forward on the head and bearing anterior orientation in life. Maxil- lary and premaxillary teeth well-developed "fangs" strongly re- curved toward tongue and embedded within the labial mucosa so that only tips of teeth protrude into buccal cavity. Dentigerous processes of prevomers exceedingly small, lying approximately equi- distant from each other and medial margins of choanae, bearing two and three fang-like teeth on right and left processes, respec- tively. Choanae large, subcircular and widely separated medially. Tongue large, circular, with posterior border free, unnotched.

Forelimb robust, lacking dennal wrist fold. First finger ( Fig. 2a) much longer than second and about equal in length to third; length of fourth finger slightly subequal to that of third; digital length in decreasing order, III-IV-I-II; phalangeal fonnula 2-2-3-3. Palmar webbing absent. Tips of fingers slightly expanded into spherical pads and lateral margins of digits with distinct fringes. Inner palmar tubercle large, elliptical and depressed. Outer palmar tubercle prominent, quadrangular and elevated. One large, round subarticular tubercle present on each of first two fingers; each of digits III and IV with two, smaller, round subarticular tubercles. Supernumerary palmar tubercles present and well developed. Hind limbs long and slender, approximately twice length of body (Table 1 ) . Toes long and slender ( Fig. 2b ) , in decreasing order of length,

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 2. — Hand and foot of Tclmatohius pefauri: a) palmar view; b) plantar view. Line equals 5 mm.

IV-III-V-II-I; phalangela fomuila 2-2-3-4-3. Webbing fonnula I 1^ - -2 II 1% -2'AUl2-3 IV 3 - 1 V; interdigital webbing smooth- ly concave, not sharply indented, diminishing distally to fringes along lateral margins of toes. Tips of toes spherical, slightly smaller than tips of fingers. Inner metatarsal tubercle small, elliptical and ele- vated. Outer metatarsal tubercle scarcely evident as an extremely small, round protuberance. Subarticular tubercles present but small

A NEW SPECIES OF TELMATOBIUS FROM CHILE 5

and poorly developed on toes; the distribution of subarticular tu- bercles per toe as follows: 1(1), 11(2), 111(2), IV(3) and V(l); supernumerary tubercles small, scattered, few in number. Distinct tarsal fold extending length of tarsus and diminishing distally into fringe on inner margin of first ( I ) toe.

Dorsal skin smooth except for few, scattered, low tubercles in presacral region; profusion of small tubercles on flanks, ventral surface of foreami, knee, outer surfaces of tibia and tarsus and posteroventral surfaces of thighs; proliferation of small skin folds extending from posterior region of head posteroventrally along flanks. One, distinct, fleshy dennal ridge fonning arc from postero- dorsal corner of axilla downward across mid-flank region. Ventral surfaces smooth except for few, low tubercles in cloacal region. Cloacal opening directed posteriorly at dorsal level of thighs; open- ing oriented transversely and inconspicuously ornamented below by folded, tuberculate skin.

Color in preservative: Unifonn drab reddish brown above. Lips, side of head, anterior surface of upper ann, flanks and anterior surfaces of thighs gray to tan with indistinct mottling and small spots of dark, reddish brown. Gular and pectoral regions and inner surfaces of forelimb immaculate grayish tan; abdominal region darker gray-tan with fine gray mottling peripherally; ventral thighs and inner surfaces of hind limbs tan with moderately bold gray mottling along posterior margins of thighs and finer gray mottling peripheral to pale areas of hind limbs. Palmar and plantar surfaces uniform dark gray-brown except for creamy tan digital tips.

Color in life: Dorsal surfaces olive-brown with distinct, darker brown spots and mottling. Ventral surfaces grayish white, and an- terior surfaces of thighs yellow-orange. Iris drab olive-brown with darker reticulations.

Chromosomes: Corneal epithelium chromosome preparations from the holotype revealed that TeJmatobiiis pefauri has a diploid number of 26 metacentric and submetacentric chromosomes.

Distribution. — This species is known only from the type locality, a small stream at the village of Mumiuntani located on the Pacific slopes of the Andean cordillera in northern Chile.

Remarks. — The single example of this distinctively long-limbed and large-headed Telmatobius was found at night underwater in a stream pool about 0.5 m deep. The stream is relatively small ( about 1.5 m wide), lacks vegetation, and is characterized by scattered, shallow pools. The stream bed is composed of sand and flat, yellow stones. Because of the terrain and local usage, the stream gradient varies from moderately rapid to slow; downstream, damming has created a small reservoir utilized by livestock. This precordilleran locality is diy and characterized by a rocky substrate with low grasses and scattered bushes less than 1 m in height. Biifo spinu-

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Table 1. — Measurements of the Holotype ( KU 159836, adult $ ) of

Telmatohius pefauri.

Character Measurement (n\m)

Snout-vent Length 75

Head Length 28

Head Width 34

Internarial Distance 5

Orbitonarial Distance 5

Diameter of Eye 8

Tibia Length 38

Tarsus Length 37

Total Foot Length 61

losus are abundant in this area, but aside from Bufo, the only other amphibian collected in several trips to this locality is the holotype of Telmatohius pefauri. Although most Telmatohius are notoriously secretive and elusive, we hope that futrn-e field work at Mur- muntani will yield adults of both sexes and larvae of this species. The female emitted a release call similar to the clucking of a hen (Veloso, pers. observation).

Discussion

Because of our limited knowledge of Tehnatohius pefauri and the confused status of the numerous (±26) and geographically widespread (Ecuador to Argentina and Chile) species included in the genus, it is prematiu-e to comment on the relationships of pefauri in more than a superficial way. This species differs from the five other known species of Tehiritohius in Chile (marmoratus, peruvi- anus, halli, montanus and laevis) in being larger and having pro- portionally longer hind limbs. Tehnatohius pefauri can be distin- guished from marmoratus bv the smoother skin (granular in the Tnarmoratus that we have observed) and paler coloration of the fonner. The shape of the head and coloration differentiate pefauri from peruvianus, which has a more acuminate snout in dorsal view and is darker in color. Tehnatohius halli differs from pefauri by the extremely depressed head and long, flat snout of the fomier. In contrast to marmoratus, peruvianus and halli, which occur in northern Chile, montanus and laevis are known only from the central Chilean cordillera, far south of the range of the northern species. Both of the southern species are characterized by moder- ately small size, short hind limbs and short, blunt snouts-a char- acter suite that readily distinguishes them from pefauri.

Vellard ( 1951 ) defined four distributional and ecological groups of Tehnatohius, as follows: 1) peruvianus group inhabiting streams and small ponds of the Andes of northern Argentina and Chile; 2) marmoratus group utilizing streams of the Titicaca Basin of Peru

A NEW SPECIES OF TELMATOBIUS FROM CHILE 7

and Bolivia; 3) jelskii group occurring in Andean streams of central Peru; and 4) ignavus group found in a variety of Andean habitats in northern Peru and Ecuador. On the basis of Vellard's (1955) analysis of the jelskii group, pcfauri would seem to be allied most closely with this group morphologically. Among the characters that are shared are the following: 1) snout shape, 2) concealed tym- panum, 3) tongue attached to the floor of the mouth over two-thirds of its length, 4) long hind limbs, 5) complete tarsal fold, 6) smooth skin, and 7) large, spherical and protuberant eyes, dorsolateral in position and directed anteriorly. However, notable differences prevail in the shape of the choanae and in the presence of pre- vomerine teeth in pefauri. Although this concordance of character states is real, we are not prepared to accept, ex facie, that pefauri should be considered a member of the jelskii group-a substantial evaluation of the systematic and evolutionary significance and variation of these and many other suites of characters is needed in order to detennine their relative value in assessing the relationships within this complex and widespread genus.

It seems appropriate at this point to comment on some of the characters mentioned above. Snout shape is a useful specific char- acter that basically reflects the osteological configuration of the maxillaiy-premaxillary arch and the orientation of the alaiy processes of the premaxillaiy. Preliminary evidence (Trueb, 1977) suggests that there is sui-prisingly little variation in snout shape within a species, although there are some instances of sexual dimorphism in frogs (e.g., see Duellman, 1970:84 and 447). Cursory perusal of anuran systematic studies indicates that this is an unreliable char- acter to utilize above the species level. Within those groups that Vellard has studied, he ( 1951 ) suggested that there is a trend toward loss of maxillary and pre vomerine dentition; the loss of this dentition is substantiated further by Capurro (1955). The trend towards loss of teeth within a genus is interesting and perhaps unique. Loss of teeth on the maxillary arch unifonnly characterizes only four anuran families-Bufonidae, Rhinodermatidae, Brachy- cephalidae and Rhinophrynidae. Most families possess some genera that are edentate (see Lynch, 1973, for a partial summary), but so far as we know, the edentate condition is consistent within the majority of genera ( Uperolia and Pliysalaemus being notable ex- ceptions). In the same paper, Vellard (1951:34) in his diagnosis of Telmatohius marmoratus stated: ''Dientes maxilares inferiores siempre presentes." We assume that "Inferiores" is a lapsus for "superiores": thus changing the meaning of the phrase to "upper maxillary teeth always present." If not, one would assume that he is referring to the presence of mandibular tecth-a condition known to occur only in the hylid frog, Amphignathodon guentheri.

The presence or absence of a tympanum has been shown to be of

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

doubtful value in Telmatohius. Vellard (1955:11-12) stated that within discrete populations of jelskii, longitarsis, rimac, walkeri and arequipensis the tympanum may be present or absent. Furthennore, in some species unifonnly lacking a tympanum, it has been found (Trueb, pers. observation) that the tympanic annulus underlying the skin may be present, absent or reduced with no apparent cor- relation with sex, age or population. Thus, the ear, as in the case of the dentition, seems to be undergoing reduction in some members of this genus.

The condition of the skin also is highly variable. Vellard (1951: 34) pointed out in his diagnosis of Telmatohius marmoratus that depending upon the "fonn," the skin may be smooth, tuberculate, warty, and with or without folds. Moreover, the condition of the skin of males of some species is known to change seasonally.

Probably two of the most characteristic features of the genus as a whole are the position and orientation of the eyes. Generally the eyes tend to be protuberant, usually dorsad on the head and fre- quently directed frontally. Unfortimately, these qualities, although obvious, are difficult to describe in such a way that interspecific comparisons are meaningful. Furthennore, it should be kept in mind that apparent differences in the size and position of the eye may be partly a function of, or exaggerated by, the shape and proportions of different parts of the head.

Before we can begin to assess the intrageneric relationships within Telmatohius, prodigious amounts of alpha taxonomic work remain to be done. Tadpoles are descrilDcd for only a few species. The osteology and the extent and natiu-e of vocalization are virtu- ally unknown. The external morphology of the hands and feet has been described only sporadically. One of us (Veloso) is work- ing with chromosomal characteristics of various Tehnatohius. In- tegration and synthesis of these kinds of infonnation with ecological, life history and distributional data should result in a better under- standing of this peculiar group of leptodactylid frogs and their rela- tionships to other members of the family.

Resumen

Se describe una nueva especie de Tehnatoliius para el Norte de Chile, Tehnatohius pefauri, sp. nov. La descripcion se basa en un ejemplar hembra adulto captiu-ado en la localidad de Mumiuntani, Departamento de Arica, Provincia de Tarapaca, Chile. Los car- acteres de la diagnosis que combinados separan esta especie de los restantes representantes del genero son los siguientes: 1) patas posteriores largas, articulacion tibiotarsal sobrepasa la orbita cuando la pata es llevada hacia adelante; 2) hocico redondeado en vista dorsal, cabeza deprimida en vista lateral; 3) arco mandibular no

A NEW SPECIES OF TELMATOBIUS FROM CHILE 9

sobrepasa la maxila; 4) narinas ligeramente protriiidas; 5) dientes maxilares y premaxilares recubiertos por pliegues mucosos; 6) procesos dentigeros pievomerianos pequeiios, orientados horizon- talmente, medianamente separados entre coanas de tamano grande; 7) dientes mandibiilaies ausentes; 8) ojos dorsolaterales proyectados hacia adelante; y 9) pliegue tarsal bien desairollado.

El tamano de la poblacion de Telmatobius pefauri se considera pequeiio, por cuanto sucesivas prospecciones del area no ban per- mitido la captura de nuevos ejemplares. De acuerdo con las descrip- ciones de Telmatobius seilaladas por otros autores esta nueva especie se relacionaria desde el punto de vista de su morfologia con especies del giiipo jelskii. Sin embargo, una sustancial evalua- cion de la sistematica y variacion de los caracteres observados es necesaria antes de detemiinar la inclusion de T. pefauri en alguna de las lineas fileticas reconocidas en Telmatobius.

Telmatobius pefauri, se reconoce facilmente de las otras cinco especies de Telmatobius seiialadas para Chile (marmoratus, peru- vianus, halli, laevis y )nontamis), por su talla y la longitud relativa de sus extremidades inferiores. T. pefauri se distingue de mar- moratus por su piel lisa y su coloracion, la fonua de la cabeza y la coloracion, diferencian pefauri de peruvianus. Telmatobius halli difiere de pefauri por su cabeza deprimida en sentido dorsoventral y por la longitud de su hocico. Telmatobius pefauri se diferencia de las especies montanus y laevis conocidas solamente para la Cordil- lera central de Chile, frente a Santiago (Lat. 33° S), ambas especies de Telmatobius del Sur estan caracterizadas por su talla moder- adamente pequefia, patas posteriores mas cortas y hocico redon- deado.

Literature Cited

Capurro, L. F. 1950. Batracios de Taiapaca. Inv. Zool. Chilenas, 1(1):9-12.

Capurro, L. F. 1953. Telmatobius marmoratus (Dum. et. Bib.) (Anura: Leptodactylidae ) nueva especie para Chile. Ibid., 2(2) : 19-22.

CAPxmRO, L. F. 1955. Telmatobius halli edentatus. Nueva subespecie para la fauna anfibia de Chile. Ibid., 2(9-10) : 150-152.

DuELLMAN, W. E. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas, ( 1 ) :.\ii -{- 753 pp.

Lynch, J. D. 1973. The transition from archaic to advanced frogs. In Evolu- tionary biology of the anurans. Contemporary research on major problems. J. L. Vial, editor. University of Missouri Press, Columbia, Missouri, pp. 133-182.

Noble, G. K. 1938. A new species of frog of the genus Telmatobius from Chile. Anier. Mus. Novitates, (973): 1-3.

Schmidt, K. P. 1954. Notes on frogs of the genus Telmatobius with descrip- tions of two new Peruvian species. Fieldiana • Zoology, 34(26) : 277-287.

Truer, L. 1977. Osteology and anuran systematics: Intiapopulational varia- tion in Hijla lancifomis. Sys. Zool., in press.

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Vellard, J. 1951. Estudios sobre batracios andinos. I. -El grupo Telmatobius y formas afines. Mem. Mus. Hist. Nat. "Javier Prado," ( 1 ) : 1-89 + 8 pp. of plates.

Vellard, J. 1955. Estudios sobre batracios andinos. III. -Los Telmatobius del grupo Jelskii. Ibid., (4): 1-28.

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