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J1ELDIANA • ZOOLOGY

Published by
CHICAGO NATURAL HISTORY MUSEUM

Volume 34 JULY 23, 1954 No. 26

NOTES ON FROGS OF THE GENUS
TELMATOBIUS

WITH DESCRIPTIONS OF TWO NEW
PERUVIAN SPECIES

KARL P. SCHMIDT

CHIEF CURATOR, DEPARTMENT OF ZOOLOGY

The aquatic Andean frogs of the genus Telmatobius have at-
tracted the attention of a succession of herpetologists on account of
their remarkable adjustment to high altitudes and to cold waters.
It adds to the interest of the group that the species adjust also to
the waters of hot springs and several species appear to be confined
to such thermal waters. Since my redescription of Telmatobius
peruvianus, the type species of the genus, in 1928, from essentially
topotypical specimens collected by my colleague Colin Campbell
Sanborn near Tacna, Peru, both he and I have continued to collect
these frogs at every opportunity. We have thus accumulated con-
siderable collections in Chicago Natural History Museum. I had
borrowed other collections in the hope of reviewing the genus, when
the appearance of an excellent and comprehensive paper by Dr. J.
Vellard (1951) on Telmatobius and allied forms makes such a review
unnecessary. Dr. Vellard has, in fact, accomplished exactly what
was needed in the genus, namely, the definition and establishment
of the two widespread species marmoratus and jelski, together with
an account of the variation of their constituent populations.

In my own preliminary studies of Peruvian Telmatobius I had
found wide differences in the size of the spines on the highly de-
veloped "thumb-pad" of the breeding male. Such secondary sex
characters have always proved to be an extremely useful key to
definitive taxonomic definition of species, and they are not much
employed by Vellard, who describes them only in general terms.
I accordingly wish to describe two new forms distinguished on this
basis, respectively from the areas assigned in Dr. Vellard's paper to

No. 733 277

THE LIBRARY OF THE

AUG 3 1954

UNIVERSITY OF ILLINOIS

278 FIELDIANA: ZOOLOGY, VOLUME 34

jelski and marmoratus. I am especially grateful to the Museum of
Zoology of the University of Michigan for the loan of a specimen
designated as the type of one of the new forms.

Telmatobius rimac, sp. nov.

Type. — University of Michigan, Museum of Zoology, no. 55596,
adult male, from the Rio Blanco, tributary of the Rio Rimac, Peru,
at an altitude of about 10,000 feet. Collected by C. H. Eigenmann
and W. R. Allen, September 18, 1918.

Diagnosis. — A frog with stocky body; skin smooth; no trace of
tympanum; vomerine teeth well developed; toes webbed to tips,
but webs emarginate; a strong fold extending backward from the
eye, expanding to form a parotoid gland; inner and outer metatarsal
tubercles well defined; spines of pad on first finger of male very fine
and closely set, eight in one millimeter.

Description of type.— A Telmatobius of moderate size, the ventral
surface yellowish; vomerine teeth very large, each group equal in
size to one of the large choanae; dorsal skin entirely smooth, ventral
skin with transverse plicae; no trace of tympanum; toes fully webbed,
webs emarginate between toe- tips; inner and outer metatarsal tu-
bercles well defined ; metatarsals partly separated ; a strong glandular
fold extending backward from the eye to above the arm, widening
posteriorly to form a parotoid gland, which, however, is not evident
from above; snout short, rounded in profile; nostrils as far apart as
their distance from the eye.

Dark brown above, without markings; dark yellowish brown be-
neath, with darker spots.

Nuptial pad on first finger minute, closely set spines, eight in
line in one millimeter; arm greatly thickened; no spines on arms
or chest.

Measurements of type. — Length 57 mm., width of head 21 mm.,
hind leg 89 mm., tibia 28 mm., arm 34 mm.

Notes on paratypes. — Three specimens (C.N.H.M. no. 34224)
collected by myself at Huinco, Rimac River, at the power dam above
Lima, September 1, 1939, are transforming or just transformed
individuals. One with a body length of 34 mm. has a stub of tail
13 mm. long. The largest specimen, measuring 38 mm., has no
trace of a tail.

Comparisons. — The Rimac Valley species is distinguished from
the widespread Telmatobius jelski because of the unmistakable dif-

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latirostns

W
%brevipes

walkeri

pustulosus** ^%'sanborm
intermedius • )

' '\ .verrucosus

marmoratus

parkeri

angustipes

culeus

escomeli

crawfordi

aemaricus

punensis

oxycephalus
hauthali

FIG. 46. Map of western South America, showing the type localities of the
species of Telmatobius (including forms in synonymy).

279

OF

280 FIELDIANA: ZOOLOGY, VOLUME 34

ference in the spines of the thumb-pad of the breeding male. There
are eight spines to the millimeter, as compared with four to the
millimeter in the large series of Telmatobius j'elski, with which the
type has been compared. Vellard refers specimens from Sta. Eulalia
in the Rimac Valley to jelski. Telmatobius rimac differs sharply
from marmoratus (which also has small spines on the thumb) in its
more fully webbed toes.

Telmatobius sanborni, sp. nov.

Type. — Chicago Natural History Museum no. 40247, adult male,
from Limbani, Puno, Peru, about 13,000 feet altitude. Collected
October 1, 1941, by Colin Campbell Sanborn.

Diagnosis. — A Telmatobius with deeply emarginate webs, dorsal
skin strongly tuberculate, a projecting lateral fold, parotoids well
developed, strong vomerine teeth in transverse rows, and with first
finger of male armed with large spines (three in two millimeters).

Description of type. — Habitus of typical Telmatobius marmoratus;
no tympanum; a laterally projecting parotoid gland, beginning as a
fold behind the eye; a post-rictal gland sharply set off by folds;
dorsal skin strongly tuberculate, with a lateral fold, below which
the sides are smooth; ventral skin granulate or plicate; vomerine
teeth in strongly developed, transverse series, projecting laterally
beyond the inner borders of the small choanae; toes with a fringe
of web to the tips, but webs very deeply emarginate, first toe essen-
tially free, as are three distal phalanges of fourth toe; an elongate
inner metatarsal tubercle, two-thirds the length of the inner toe;
outer metatarsal tubercle small, round, distinct.

Dark gray above, lighter beneath.

Arm enormously hypertrophied, the inner finger bearing a raised
pad covered with large spines, which measure about 0.66 mm. in
diameter at the base, that is, three spines to two millimeters; scat-
tered spines on arms and breast.

Measurements of type. — Length of body 60 mm.; width of head
at angles of jaws 27 mm.; hind leg 96 mm.; tibia 32 mm.; (arm not
extendable), diameter of arm at elbow 13 mm.

Notes on paratypes.—Five males (nos. 67435-67438 and 40248a),
the largest measuring 65 mm. from snout to anus, correspond with
the description of the type. The nine female specimens (no. 40248)
range in size from 26 mm. to 48 mm. In the largest specimen the
hind leg measures 83 mm., tibia 26 mm., arm 32 mm., width of head

SCHMIDT: FROGS OF GENUS TELMATOBIUS 281

18 mm. In some of these specimens the dorsal tubercles are seen
to correspond to black spots, and the inner faces of thigh and tibia
may be yellow.

Comparisons. — While this species is most naturally to be com-
pared with Telmatobius marmoratus, it appears to be much more
distinct from m. marmoratus of the Lake Titicaca region than are
m. pustulosus of Cuzco or m. hintoni of Bolivia. Limbani is in the
drainage of the Rio Inambari, and the range of the new form pre-
sumably extends downward from that locality to the limits of cold
water. Telmatobius sanborni differs from all other species examined
in the larger size of the tubercles on the first finger of the breeding
male.

Telmatobius escomeli exsul Vellard

Telmatobius escomeli exsul Vellard, 1951, Mem. Mus. Hist. Nat. "Javier
Prado," 1 : 66, fig. 19 — Pozo de agua junto a las termas de Yura, cerca
de Arequipa, 2,575 meters.

Vellard records specimens of Telmatobius marmoratus from the
vicinity of the hot springs of Jesus, from a warm spring at Arequipa,
and from a "pozo de agua tibia" at the hot springs of Yura. Speci-
mens from the Arequipa region in Chicago Natural History Museum
collections average much larger than the marmoratus recorded by
Vellard, and thus require comparison with exsul. They are quite
evidently not marmoratus.

CHECK LIST OF SPECIES

Boulenger recognized six species of Telmatobius in 1882, and by
1920 this had risen to thirteen, after the exclusion of the species
of Cyclorhamphus (Barbour and Noble, 1920). Parker (1940) con-
firms the suggestion of Barbour and Noble that Telmatobius mar-
moratus Dume'ril and Bibron is from Peru, not Chile; he greatly
clarifies the ecological relations of the species characterized by an
extremely baggy skin, which appears especially in the lacustrine
forms; and he describes five new forms. Vellard finally (1951)
recognizes nineteen species, of which four are divided into subspecies,
making twenty-five forms in all. Following Vellard's classification
closely, except as to the sub-specific arrangement of albiventris,
with a few additional references, the following list results.

It is hoped that the arrangement of the species according to
the north-south latitudes of their type localities, which are shown

282 FIELDIANA: ZOOLOGY, VOLUME 34

on the accompanying map, may be suggestive for further subspecific
reclassification.

Telmatobius Wiegmann

Telmatobius Wiegmann, 1835, Nova Acta Acad. Leop. Carol., 17: 262 —

type: peruvianus.
Pseudobatrachus Peters, 1873, Monatsber. Akad. Wiss. Berlin, 1873: 414 —

type: jelskii.

Telmatobius niger Barbour and Noble

Telmatobius niger Barbour and Noble, 1920, Bull. Mus. Comp. Zool., 63:

413, fig. 4 — Palmira desert, 10,500 feet altitude, Ecuador (=vicinity of
Palmira, Chimborazo Province).

Telmatobius cinereus Noble

Telmatobius cinereus Noble, 1921, Amer. Mus. Nov., 29: 6 — Bestion [Azuay],
Ecuador.

Telmatobius ignavus Barbour and Noble

Telmatobius ignavus Barbour and Noble, 1920, Bull. Mus. Comp. Zool., 63:

414, fig. 5 — Huancabamba, Piura, Peru.

Telmatobius latirostris Vellard

Telmatobius latirostris Vellard, 1951, Mem. Mus. Hist. Nat. "Javier Prado,"
1 : 68, fig. 22— Cutervo, Cajamarca, Peru.

Telmatobius brevipes Vellard

Telmatobius brevipes Vellard, 1951, Mem. Mus. Hist. Nat. "Javier Prado,"
1: 71, figs. 20-21— Huamachuco, Libertad, Peru.

Telmatobius jelski Peters

Pseudobatrachus jelskii Peters, 1873, Monatsber. Akad. Wiss. Berlin, 1873:

415, pi. 4, fig. 1 — Acancocha, Junin, Peru.

Telmatobius jelskii Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., p. 191.

Telmatobius rimac, sp. nov.
See page 278.

Telmatobius walkeri Shreve

Telmatobius walkeri Shreve, 1941, Proc. New England Zool. Club, 18: 79 —
Rapi, 108 km. east of Ayacucho, Ayacucho, Peru.

SCHMIDT: FROGS OF GENUS TELMATOBIUS 283

Telmatobius sanborni, sp. nov.

See page 280.

Telmatobius intermedius Vellard

Telmatobius intermedius Vellard, 1951, Mem. Mus. Hist. Nat. "Javier Prado,"
1 : 29, fig. 2 — Allipacca, near Puquio, Ayacucho, Peru.

Telmatobius marmoratus BiimeYi! and Bibron

Cyclorhamphus marmoratus Dumeril and Bibron, 1841, Erpet. Gen., 8: 455
— Guasacona, Chile (=Guasacoma, Puno, Peru).

Telmatobius marmoratus Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., p. 192.
Cyclorhamphus aemaricus Cope, 1874, Proc. Acad. Nat. Sci. Phila., 125 —
Lake Titicaca, Puno, Peru.

Cyclorhamphus angustipes Cope, 1877, Proc. Amer. Phil. Soc., 17: 38 —
Juliaca, Puno, Peru.

Telmatobius marmoratus marmoratus Dumeril and Bibron

Telmatobius marmoratus marmoratus Parker, 1940, Trans. Linn. Soc. London,
(3), 1:208, fig. 1.

Telmatobius marmoratus pustulosus Cope

Cyclorhamphus pustulosus Cope, 1877, Proc. Amer. Phil. Soc., 17: 39 — Tinta,
Cuzco, Peru, 11,400 feet altitude.

Telmatobius marmoratus pustulosus Parker, 1940, Trans. Linn. Soc. London,
(3), 1:208, fig. 2.

Telmatobius marmoratus hintoni Parker

Telmatobius hintoni Parker, 1940, Trans. Linn. Soc. London, (3), 1: 209,
• fig. 3— Tolota, Cochabamba, Bolivia (=Tolata).

Telmatobius marmoratus hintoni Vellard, 1951, Mem. Mus. Hist. Nat. "Javier
Prado," 1:38.

Telmatobius culeus Garman

Cyclorhamphus culeus Garman, 1875, Bull. Mus. Comp. Zool., 3: 276, pi.
— Lake Titicaca, off Achacache, 11 fathoms depth.

Telmatobius culeus Barbour and Noble, 1920, Bull. Mus. Comp. Zool., 63: 416.

Telmatobius culeus culeus Garman

Telmatobius culeus culeus Parker, 1940, Trans. Linn. Soc. London, (3), 1:
210, fig. 5.

284 FIELDIANA: ZOOLOGY, VOLUME 34

Telmatobius culeus albiventris Parker

Telmatobius culeus albiventris Parker, 1940, Trans. Linn. Soc. London, (3),
211, fig. 6 — Lago Pequeno, Lake Titicaca (near Taquiri Island and
Taraco Point).

Telmatobius culeus punensis Vellard

Telmatobius albiventris punensis Vellard, 1951, Mem. Mus. Hist. Nat. "Javier
Prado," 1: 57 — Isla Amantani, opposite the town of Puno, Puno, Peru.

Telmatobius culeus parkeri Vellard

Telmatobius albiventris parkeri Vellard, 1951, Mem. Mus. Hist. Nat. "Javier
Prado," 1 : 58, fig. 15 — Lake Arapa, north of Lake Titicaca.

Telmatobius escomeli Angel

Telmatobius escomeli Angel, 1923, Ann. Sci. Nat. (zool.), 6: 108 — "Lake
Titicaca" (corrected to Lagunilla Lagunilla, Puno, Peru).

Telmatobius escomeli escomeli Angel

Telmatobius escomeli escomeli Parker, 1940, Trans. Linn. Soc. London, (3),
1:212, fig. 7.

Telmatobius escomeli crawfordi Parker

Telmatobius escomeli crawfordi Parker, 1940, Trans. Linn. Soc. London, (3),
1 : 212, fig. 8 — Lagunilla Saracocha, Puna, Peru.

Telmatobius escomeli exsul Vellard

Telmatobius escomeli exsul Vellard, 1951, Mem. Mus. Hist. Nat. "Javier
Prado," 1 : 66, fig. 19 — pond near the hot springs at Yura, Arequipa, Peru.

Telmatobius verrucosus Werner

Telmatobius verrucosus Werner, 1899, Zool. Anz., 22: 482 — Chaco, Bolivia
(=Chaco, on the Unduavi River, 25 km. northwest of Puente de la Via),
La Paz, Bolivia.

Telmatobius bolivianus Parker, 1940, Trans. Linn. Soc. London, (3), 1: 213,
fig. 9 — same locality as for verrucosus.

Telmatobius peruvianus Wiegmann

Telmatobius peruvianus Wiegmann, 1835, Nova Acta Acad. Leop. Carol., 17:
262, pi. 22, fig. 2— Peru (Cordillera de Guatilla, near Palca, two days'
journey east of Tacna).

SCHMIDT: FROGS OF GENUS TELMATOBIUS 285

Telmatobius simonsi Parker

Telmatobius simonsi Parker, 1940, Trans. Linn. Soc. London, (3), 1: 210,
fig. 4 — Sucre, Chuquisaca, Bolivia.

Telmatobius halli Noble

Telmatobius halli Noble, 1938, Amer. Mus. Nov., 963: 1— Ollague, Anto-
fagasta, Chile, 10,000 feet altitude.

Telmatobius oxycephalus Vellard

Telmatobius oxycephalus Vellard, 1946, Acta Zool. Lilloana, 3: 320 — Cerro de
Escalera, Salta, Argentina, 3,800 meters altitude.

Telmatobius hauthali Koslowsky

Telmatobius hauthali Koslowsky, 1895, Rev. Mus. La Plata, 6: 359, pi. 1 —
Aguas Calientes, near Cazadero Grande, Catamarca Argentina, 4,000
meters altitude.

Telmatobius hauthali hauthali Koslowsky

Telmatobius hauthali hauthali Vellard, 1951, Mem. Mus. Hist. Nat. "Javier
Prado," 1 : 79.

Telmatobius hauthali schreiteri Vellard

Telmatobius schreiteri Vellard, 1946, Acta Zool. Lilloana, 3: 321 — Aimogasta,

La Rioja, Argentina.
Telmatobius hauthali schreiteri Vellard, 1951, Mem. Mus. Hist. Nat. "Javier

Prado," 1: 80.

Telmatobius laevis Philippi

Telmatobius laevis Philippi, 1902, Suppl. Batr. Chilenos, p. 4 — Potrero, Chile
(=Potrero Grande, Province of Aconcagua, cf. Miiller, 1938).

Telmatobius montanus Philippi

Telmatobius montanus Philippi, 1902, Suppl. Batr. Chilenos, p. 47 — lake in
the high Andes, Province of Santiago, Chile (believed by Miiller, 1938, to
be near Potrero Grande, Province of Aconcagua).

REFERENCES

ANGEL, FERNAND

1923. Surun batracien nouveau du Perou appartenant au genre "Telmatobius."
Ann. Sci. Nat. (bot. zool.), 6: 107-111, fig. Describes Telmatobius escomeli
as new.

286 FIELDIANA: ZOOLOGY, VOLUME 34

BARBOUR, THOMAS, and NOBLE, G. K.

1920. Some amphibians from northwestern Peru, with a revision of the genera
Phyllobates and Telmatobius. Bull. Mus. Comp. Zool., 63: 393-427, pis. 1-3,
text figs. 1-8. Revision of Telmatobius, describing T. ignavus and niger
as new.

BOULENGER, G. A.

1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of
the British Museum. London, British Museum, xvi+503 pp., 30 pis., 49
text figs. Summarizes the genus Telmatobius, pp. 190-193.

COPE, E. D.

1874. On some Batrachia and Nematognathi brought from the Upper Amazon

by Prof. Orton. Proc. Acad. Nat. Sci. Phila., 26: 120-137. Describes

Cyclorhamphus aemaricus from Lake Titicaca.
1877. Synopsis of the cold-blooded Vertebrata, procured by Prof. James

Orton during his exploration of Peru in 1876-77. Proc. Amer..Phil. Soc., 17:

33-49. Describes Cyclorhamphus angustipes and C. pustulosus, with notes

on C. aemaricus.

DUMERIL, A. M. C., and BIBRON, GABRIEL

1841. [Tome huitieme], comprenant 1'histoire general des batraciens, et la
description des cinquante-deux genres et cent soixant-trois especes des deux
premiers sous-ordres: les peromeles qui n'ont pas de membres, et des anoures
qui sont prives de la queue. Erpet. Gen., 8: iii+792. Describes Telmatobius
marmoratus (as Cyclorhamphus m.), p. 455.

GARMAN, SAMUEL

1876. Exploration of Lake Titicaca, by Alexander Agassiz and S. W. Garman.
I. Fishes and Reptiles. Bull. Mus. Comp. Zool., 3: 273-278, pi. Describes
as new Telmatobius culeus from Lake Titicaca (as Cyclorhamphus), with notes
on Telmatobius marmoratus.

KOSLOWSKY, JULIO

1895. Batracios y reptiles de Rioja y Catamarca (Republica Argentina)
recogidos durante los meses de Febrero a Mayo de 1895 (expedition del
director del Museo). Rev. Mus. La Plata, 6: 357-370, pis. 1-4. Describes
and figures Telmatobius hauthali as new, from Catamarca.

MtJLLER, LORENZ

1938. Beitrage zur Kenntnis der Herpetofauna Chiles. X. Uber ein exemplar
von Telmatobius montanus Philippi. Zool. Anz., 121: 313-317, figs. 1-2.
Redescribes Telmatobius montanus from breeding male, and establishes a
type locality.

NOBLE, G. K.

1921. Five new species of Salientia from South America. Amer. Mus. Nov.,
29: 1-7, figs. 1-6. Describes Telmatobius cinereus.

1938. A new species of frog of the genus Telmatobius from Chile. Amer.
Mus. Nov., 973: 1-3. Describes Telmatobius halli.

PARKER, H. W.

1940. Percy Sladen Trust Expedition to Lake Titicaca in 1937 under the leader-
ship of Mr. H. Gary Gilson, M. A. xii. Amphibia. Trans. Linn. Spc. London,
(3), 1 : 203-216, 9 figs. Reviews the form of the Lake Titicaca region, describ-
ing as new Telmatobius hintoni, simonsi, bolivianus, escomeli crawfordi, and
culeus albiventris.

PETERS, WILHELM

1873. Uber neue oder wenig bekannte Gattungen und Arten von Batrachiern.
Monatsber. Akad. Wiss. Berlin, 1873: 411-418, pis. 1-4. Describes Bat-
rachophrynus with B. macrostomus and B. brachydactylus, and Pseudobat-
rachus, with P. jelski (=Telmatobius jelski).

SCHMIDT: FROGS OF GENUS TELMATOBIUS 287

PHILIPPI, R. A.

1902. Supplemento a los batraquios chilenos descritos en la Historia Fisica i
Politica de Chile de Don Claudio Gay. Santiago de Chile, Libreria Alemana
de Jose Ivens: 1-161. Describes Telmatobius laevis and montanus.

SCHMIDT, K. P.

1928. The Chilean frogs of the genus Telmatobius. Rev. Chilena Hist. Nat.,
32: 98-105. Redescribes Telmatobius peruvianus, T. laevis, and T. montanus.

SHREVE, BENJAMIN

1941. Notes on Ecuadorian and Peruvian reptiles and amphibians with de-
scription of new forms. Proc. New England Zool. Club, 18: 71-83. Discusses
Telmatobius jelski and describes T. walkeri as new.

VELLARD, J.

1946. El genero "Telmatobius" en la Republica Argentina. Acta Zool. Lil-
loana, 3: 313-326. Redescribes Telmatobius hauthali and describes as new
T. schreiteri and oxycephahis.

1951. Estudios sobre batracios andinos. I. El grupo Telmatobius y formas
afines. Mem. Mus. Hist. Nat. "Javier Prado," 1: 1-89, figs. 1-30. Review
of the genera Batrachophrynus and Telmatobius, with examination of the
variation in T. marmoratus and T. jelski, and description of the new forms
Telmatobius latirostris, brevipes, intermedius, oxycephalus, albiventris parkeri,
a. punensis, and escomeli exsul.

WERNER, FRANZ

1899. Beschreibung neuer Reptilien und Batrachier. Zool. Anz., 22: 479-484.
Describes Telmatobius verrucosus, from "Chaco, Bolivia," as new.

WlEGMANN, A. F. A.

1835. Beitrage zur Zoologie gesammelt auf einer Reise um die Erde, von Dr.
F. J. F. Meyen, M. d. A. d. N. Siebente Abhandlung. Amphibien. Nova
Acta Acad. Leop. Carol., 17: 183-268, 268a-d, 10 pis. Describes Telmatobius,
with the single species peruvianus.

UNIVERSITY OF ILLINOIS-URBANA