Herpetologists' League

A New Frog Species of the Eleutherodactylus fitzingeri Group from the Pacific Andean
Versant in Ecuador
Author(s): John D. Lynch
Reviewed work(s):
Source: Herpetologica, Vol. 35, No. 3 (Sep., 1979), pp. 228-233
Published by: Herpetologists' League
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228 HERPETOLOGICA [Vol. 35, No. 3

LITERATURECITED GOELLNER, R. 1975. Garter snakes as toxic as

rattler. St. Louis Herpetol. Soc. Newsl. 2:8-9.
ANTHONY, J. 1955. Essai sur l'evolution anato- KLAUBER, L. M. 1939. A statistical study of the
mique de l'appareilvenimeux des ophidiens. Ann. rattlesnakes. VI. Fangs. Occas. Pap. San Diego
Sci. Nat. Zool. Biol. Anim. 17:7-53. Soc. Nat. Hist. 5:1-61.
BOGERT, C. M. 1943. Dentitional phenomena in 1956. Rattlesnakes. Their habits, life
cobras and other elapids, with notes on the histories, and influence on mankind. Vol. 1 and
adaptive modifications of their fangs. Bull. Am. 2. Univ. Calif. Press, Berkeley and Los Angeles.
Mus. Nat. Hist. 81:285-360. PEYER, B. 1963. Die Zahne, ihr Ursprung, ihre
BRATTsTRoM,B. H. 1964. Evolution of the pit Geschichte und ihre Aufgabe. Verstaendliche
vipers. Trans. San Diego Soc. Nat. Hist. 13: Wiss. no. 79, Jul. Springer, Berlin.
185-268. . 1968. Comparativeodontology. Ed. and
EDMUND, A. G. 1960. Tooth replacement phe- tr. by R. Zangerl. Univ. Chicago Press, Chicago
nomena in the lower vertebrates. R. Ont. Mus. and London.
Life Sci. Contrib. 52:1-42. TAUB, A. M. 1965. Comparative histological
. 1969. Dentition, pp. 117-200. In C. studies on the parotid gland of colubrid snakes.
Gans, A. Bellairs, and T. S. Parsons (Eds.), Ph.D. Dissertation. State Univ. New York at
Biology of the Reptilia. Vol. 1. Academic Press, Buffalo.
New York.
FRAZZETTA, T. H. 1966. Studies on the mor- Accepted: 29 October 1978
phology and function of the skull in the Boidae Department of Zoology and Electron
(Serpentes). Part II. Morphology and function
of the jaw apparatusin Python sebae and Python Microscope Center, Washington State Uni-
molurus. J. Morphol. 118:217-296. versity, Pullman, Washington 99164, USA

35(3), 1979, 228-233

Herpetologica,
? 1979 by The Herpetologists' League

A NEW FROG SPECIES OF THE ELEUTHERODACTYLUS

FITZINGERI GROUP FROM THE PACIFIC ANDEAN
VERSANT IN ECUADOR
JOHN D. LYNCH

ABSTRACT: Eleutherodactylusactites sp. nov. is a large species of the essentiallylowland

fitzingerigroupfoundin the vicinityof Pilalo,ProvinciaCotopaxi,Ecuador,at 2486 m. Eleu-
therodactylusactitesis most similarto and closelyrelatedto the allopatricE. w-nigrumdis-
tributedthroughthe Andesof Colombiaand Ecuador.UnlikeE. w-nigrum,E. actitesis most
commonin nonforestedmicrohabitats.
Key words: Amphibia;Salientia;Leptodactylidae;
Eleutherodactylus;
sp. nov.; Ecuador

LEPTODACTYLID frogs of the Eleuthero- de Santa Marta, Colombia), E. lymani Bar-

dactylus fitzingeri group are primarily low- bour and Noble (Huancabamba depression
land frogs found in forested environments in southern Ecuador and northern Peru),
(Lynch, 1976). Many species in trans- E. thectopternus Lynch (Cordilleras Cen-
Andean South America and in Middle tral and Occidental in Colombia), E. viridi-
America are closely associated with streams cans Lynch (Cordillera Occidental, Colom-
(Savage, 1974, 1975). South American bia), and E. w-nigrum (Boettger) (all three
species normally found above 1000 m in- cordilleras in Colombia, both Amazonian
clude E. insignitus Ruthven (Sierra Nevada and Pacific Andean versants in Ecuador).
September 1979] HERPETOLOGICA 229

Eleutherodactylusw-nigrum is the most

widely distributed Andean Eleutherodac-
tylus, ranging 70 N-400 S (Fig. 1) over an
altitudinal range from 800 m (Pacific ver-
sant in Colombia and Ecuador) to 3200 m
(the type-locality, near Cuenca, Ecuador)
although it seldom is found below 1400 m
or above 2800 m. Geographic variation is
apparent in coloration and color pattern
-4 / 'f,
and to a lesser extent in size at maturity
-0~~~~~
(Table 1).
Cochran and Goin (1970) provided a 0)
detailed description for this frog which can
be described as large (adult males 25.0-
46.8 mm, adult females 43.2-71.5 mm SVL), '' S @1.s
with large black spots on the flanks, spots
or marbling in the groin and on the anterior 4000km
and posterior surfaces of the thighs, diffuse
gray spotting on the venter, and white or
yellow ground color on concealed surfaces
of the limbs and flanks. Adult males have
vocal slits and non-spinous nuptial pads.
In 1968 during survey work on the Pacific
versant in Ecuador, I collected at Pilalo,
Provincia Cotopaxi (2486 m). Specimens
of a large member of the fitzingeri group FIG. l.- Map of distributions of Eleutherodac-
resembling but obviously different from E. tylus actites (square) and E. w-nigrum (circles).
The recordfor E. w-nigrumfrom the SierraNevada
w-nigrum were abundant along the head- de Santa Marta in northern Colombia is not in-
water streams of the Rio Pilalo in sparsely cluded.
wooded pastures. Examination of the major
museum collections and further fieldwork
in western Ecuador confirmed the existence The following abbreviations are used
of an undescribed species of the fitzingeri in the account below: SVL (snout-vent
group found only in the immediate vicinity length), HW (head width), IOD (inter-
of Pilalo where it apparently replaces the orbital distance), and E-N (eye to nostril
widespread E. w-nigrum. distance) .

TABLE 1.-Size at maturity of Eleutherodactylusw-nigrum in Colombia and Ecuador.

Locality Males Females

Colombiae 25.0-37.9(31.6 ? 1.3)252 44.6-56.2(50.5 ? 2.0)14

N Pacific slopes, Ecuador3 25.0-46.1(33.0 + 1.3)58 50.2-71.5(59.8 ? 2.5)21
vic. Guaranda, Ecuador' 29.1-32.6(31.0)3 43.2-46.1(44.9)3
vic. Pallatanga, Ecuador4 29.0-38.1(34.3)5 47.8-69.7(55.8)3
Azuay Prov., Ecuador' 28.7-30.6(29.6)2 45.5-57.0(51.2)2
El Oro Prov., Ecuador' 60.0-63.6(61.8)2
N Amazonian slopes, Ecuador5 29.3-46.8(34.3 ? 1.2)32 44.4-56.6(49.9 ? 1.9)16
S Amazonian slopes, Ecuador 28.4-45.1(36.7 ? 1.0)55 53.0-65.4(59.2 + 2.6)9
'Deptos. Antioquia, Cauca, Huila, Tolima, and Valle.
2 Range in mm (x ? 2 SE), n.
2 South to the Rio Toachi.
4 All localities on Pacific slopes.
5 North of the Pastaza trench.
230 HERPETOLOGICA [Vol. 35, No. 3

Eleutherodactylus actites sp. nov.

Holotype.-KU (University of Kansas
Museum of Natural History) 120111, a
gravid female, one of a series collected at
Pilalo, Provincia Cotopaxi, Ecuador, 2486
m, on 25 June 1968 by Robert W. Hender-
son and John D. Lynch.
Paratypes.-KU 120112-124, 131210-260,
141776-93; collected at type-locality on 25
June 1968, 3-4 July 1970, and 7 July 1971,
respectively.
Diagnosis.-(1) Skin of dorsum sha-
greened with thin dorsolateral folds, that A A
of venter smooth; (2) tympanum distinct,
its length 1/3-1/2 that of eye; (3) snout acu-
minate in dorsal view, rounded in lateral
profile; canthus rostralis sharp; (4) upper
eyelid as broad as IOD in males, slightly
narrower than in females, not bearing pun-
gent warts; no cranial crests; (5) vomerine
odontophores prominent, triangular in out-
line; (6) males with vocal slits, subgular
vocal sac; non-spinous nuptial pads on
thumbs; (7) first finger longer than second;
pads on fingers II-IV moderate-sized, none FIG. 2.-( A) Eleutherodactylus actites (KU
on thumb; discs broader than long; (8) 120111, holotype) and (B) Eleutherodactylus w-
fingers bearing lateral keels; (9) no ulnar nigrum (adult female, not preserved,from Tandapi,
Provincia Pichincha, Ecuador, 1460 m).
tubercles except for small antebrachial
tubercle; (10) small tubercles on heel; no
outer tarsal tubercles; inner tarsal fold along
distal 2/5 of tarsus; (11) two metatarsal E. actites requires comparison with E. acha-
tubercles, inner elongate, 4-6 times size of tinus (Boulenger), E. conspicillatus (GGun-
indistinct outer; 2-3 indistinct supernumer- ther), E. fenestratus (Steindachner), E.
ary plantar tubercles; (12) toes bearing gutturalis Hoogmoed, Lynch, and Lescure,
lateral keels, not webbed; toe pads as large E. peruvianus (Melin), E. terraebolivaris
as those of outer fingers; (13) gray with Rivero, E. vilarsi (Melin), E. w-nigrum
darker markings, viz., interorbital bar, scap- (Boettger), and E. zeuctotylus Lynch and
ular W, chevrons, labial bars; groin and Hoogmoed. E. actites differs from all of
posterior thigh speckled with black; venter these in having a short tarsal fold. Eleu-
cream spotted with gray; undersides of therodactylus actites is larger than any of
limbs dull gray; (14) adults large, 32 males the lowland species (achatinus, conspicil-
30.0-40.0 (x = 35.0) mm, 21 females 48.2- latus, fenestratus, gutturalis, peruvianus,
64.2 ( =54.9) mm SVL. terraebolivaris, vilarsi, and zeuctotylus),
On the bases of traits 1, 2, and 7, E. most of which have uniformly brown pos-
actites (Fig. 2) is referable to the fitzingeri terior surfaces of the thighs (all except
group of Eleutherodactylus (Lynch, 1976). conspicillatus and peruvianus).
In lacking webbing on the toes and having Eleutherodactylus actites is distinguished
lateral keels on the digits (as opposed to from E. w-nigrum in having an inner tarsal
distinct fringes, see Savage, 1975) and nup- fold (no fold in E. w-nigrum) and from
tial pads on the thumbs of breeding males, E. lymani in having lateral keels on the
September 1979] HERPETOLOGICA 231

digits (instead of distinct fringes). Neither %, fold not flap-like; inner metatarsal tubercle
of these close relatives has dense black elongate (three times as long as wide), non-com-
flecking on the anterior and posterior sur- pressed; outer metatarsal tubercle indistinct, sub-
conical, 1/4A/6 size of inner; indistinct supernumer-
faces of the thighs as does E. actites. ary plantar tubercles at base of toes 2, 3, and 4;
subarticular tubercles non-conical, those on inner
Description.-Head narrowerthan to as broad as toes round, those on outer toes longer than wide;
body; head broader than long; head width 33.0- toes bearing narrow, keel-like lateral fringes, not
37.6 (x = 35.4 + 0.4 [x + 2 SE], n = 71) percent webbed; all toes bearing broad pads and discs
SVL; snout acuminate in dorsal view, rounded to (broader than long), all apically rounded; hind-
weakly protruding in lateral profile, snout long, limbs long, heels broadly overlap when flexed legs
E-N 86.9-111.8 (i = 99.6 + 2.1, n = 33) percent
held at right angles to body; heel of adpressed leg
eye length in males, 105.0-134.6 (x 115.4 + 2.1, extending beyond tip of snout; shank 51.5-63.8
n = 38) percent in females; nostrils not protuber-
ant, directed dorsolaterally;canthus rostralissharp, (x = 58.4 + 1.0, n = 33) percent SVL in males,
53.7-64.7 (i = 59.5 + 1.3, n = 21) percent in
straight or weakly convex; loreal region flat, sloping
adult females, 57.8-69.0 (x = 62.1 + 1.1, n = 17)
abruptly to lips; lips not flared; upper eyelid width percent in juvenile and young females.
79.4-133.3 (x = 103.1 + 4.6, n = 30) percent
In preservative, gray above with darker gray
IOD in males, 75.0-106.2 (i = 90.5 -+- 3.1, n =
36) percent in females; interorbital space flat, no interorbitalbar, scapular W, one to three chevrons,
cranial crests; supratympanicfold prominent, con- oblique limb bars (narrower than interspaces);
cealing uppermost edge of tympanum; tympanum some individuals have numerous smaller gray spots
prominent, slightly higher than long, its length interspersed among those listed above; canthal,
(x = 38.8
supratympanic, labial stripes darker gray; some
32.2-45.7 -+- 1.2, n - 33) percent eye
individualshave labial bars; anal triangle dark gray;
length in males, 33.3-50.5 (x 42.7 ? 1.2, n = groin and posterior flanks, anterior and posterior
38) percent in females; tympanum separated from
surfaces of arm and thigh speckled with black;
eye by twice its length; no enlarged tubercles on
head except for prominent postrictal tubercles; posterior thigh dark gray with some black flecks
choanae moderate-sized, triangular in outline, not along upper edge (at posterioredge of thigh bars);
concealed by palatal shelf of maxillaryarch; vomer- venter dull cream spotted with gray; lower surfaces
ine odontophores prominent, four times size of a of limbs, bands, and feet dull gray.
choana, median and posteriorto choanae, separated In life, gray, pale reddish-brown,to dark brown
above with darker brown markings edged with
by a choanal width; odontophores approximately
triangular in outline, each bearing a row of 4-7 black; canthal and supratympanic stripes black
teeth in a nearly transverserow (dipping medially) edged with yellow-bronze;flanks paler than dorsum
across posterior edge; tongue slightly longer than with pale rose wash becoming pale blue-gray ven-
wide, posterior border shallowly notched, posterior trally and posteriorly,spotted with black; posterior
thighs pale blue-gray speckled with silver and
/4 not adherent to floor of mouth; males with long black; venter white to yellow, speckled with brown;
vocal slits posterolateral to tongue; median vocal
sac present. undersidesof thighs flesh-colored;palms, soles, and
tarsi black; iris pale yellow reticulated with black
Skin of dorsum, head, flanks, and limbs sha-
greened (no enlarged tubercles); thin, non-glan- and bearing red horizontal streak.
Measurements of the holotype in millimeters.-
dular dorsolateral folds present; skin of venter
SVL 52.2; shank 30.3; head width 18.2; head
smooth to wrinkled,some suggestion of granulations
length 19.1; tympanum length 2.2; eye length 5.6;
on posterior part of belly (anterior to discoidal
E-N 6.0. The holotype is an adult female having
fold); undersides of limbs smooth except thighs
extensively convoluted oviducts; most ovarian eggs
posteroventralto anus (coarsely areolate compared
to that of venter); anal opening not modified, no are small and white but some are large and yellow.
para-anal tubercles; no ulnar tubercles except Ety mology.-Greek, aktites, a shore
rounded antebrachial tubercle; palmar tubercle dweller, in reference to the abundance of
bifid, twice as large as oval thenar tubercle; indis-
tinct supernumerarytubercles at base of fingers 3 the frogs along edges of small streams flow-
and 4; subarticular tubercles on inner digits sub- ing through Pilalo.
conical, those on outer digits flat, as long as wide, Distribution.-Known only from the im-
simple; fingers bearing narrow, keel-like lateral mediate vicinity of Pilalo, Provincia Coto-
fringes; moderate-sized pads on fingers 2-4, none
on thumb; all digits bearing discs, that on thumb paxi, Ecuador, 2486 m. In addition to the
round, others broader than long; pads and discs large series at the University of Kansas,
rounded apically; thumb longer than second finger. there is a large uncatalogued series at the
Knee lacking tubercles; heel bearing small round
tubercles; no tubercles on outer edge of tarsus; National Museum of Natural History and
inner edge of tarsus bearing distinct fold on distal another in my possession (to be used for
232 HERPETOLOGICA [Vol. 35, No. 3

studies on reproductionand feeding) to be occurred because E. w-nigrum and E. in-

deposited at some museum. signitus are easily distinguished and Ruth-
Remarks.-One of the reasons for col- ven was familiar with each species but did
lecting at Pilalo was to gather additional not report E. w-nigrum from the Santa
altitudinal and geographical distributional Martas.
data for the frogs of the Pacific versant of Ecological observationis.-In late June
Ecuador. When I first saw E. actites I 1977, efforts were made to determine the
thought it was a geographic variant of E. habitat preferences of E. actites at Pilalo.
w-nigrum but subsequent collecting failed Three hundred and three individuals were
to yield specimens of intermediatecharac- captured and marked in a pasture at the
teristics in geographicallyproximatelocal- northwest edge of Pilalo along small streams
ities. Additional fieldwork strongly sug- (<1.0 m wide); 277 were caught by day
gested that E. w-nigrumis not part of the and 12 at night in grass. An additional 14
fauna in the vicinity of Pilalo, where it were collected at night on branches and
apparently is ecologically and geographi- vegetation (ferns and Gunnera) 0.2-1.5 m
cally replaced by E. actites. above ground. At the southeast edge of
Eleutherodactylus actites apparently is Pilalo is a larger stream (21/2 m wide) with
closely related to E. lymani Barbour and little grass or herbaceous vegetation along
Noble and E. tv-nigrum(Boettger). Aside it. Its borders consist of large gravel and
from color differences and similarities,E. boulders. Juvenile frogs were conspicuous
actites is like E. lymani in having an inner in the daytime and adults (males and fe-
tarsal fold (absent in E. w-nigrum). Eleu- males) were found beneath rocks by day.
therodactyluslymaniis readilydistinguished At night, the adult frogs were conspicuous
from E. actites in having a broader head as they sat on rocks or branches along the
(1HW40-45%SVL), weakly flaredlips, and streams. Juvenile frogs (<30 mm) were
larger tympana (tympanum length 1/2-4 found beneath rocks. One hundred and
eye length). The posteriorsurfaces of the ninety-nine of the marked frogs were dis-
thighs of E. lymani are black with white tinctly juveniles (12-27 mm SVL) and all
spots or reticulation(in life). but two were caught by day in grass. Frogs
Eleutherodactylusw-nigrum (Fig. 2) ap- identified as males range from 29-38 mm in
parently is the most widely distributed SVL and six of 46 were collected at night.
Andean eleutherodactyline ranging from Frogs identified as females range from 29-
Departamento Antioquia in Colombia 63 mm in SVL and 19 of 57 were collected
(Cochran and Goin, 1970) to Provincia at night. If females 48 mm and larger are
Zamora-Chinchipe in Ecuador. The species mature, only 12 were marked and only five
is found on the Amazonian and Pacific of those were found during the day. No
versants of the EcuadorianAndes as well calling was heard that could be attributed
as in at least two interandeanhoyas (Ota- to E. actites.
valo and Cuenca). Colombian specimens In January 1978 I returned to Pilalo and
have been found on the Pacific slopes of caught only one marked frog. Juveniles
the CordilleraOccidental, on the slope of (27 mm SVL or less) had been conspicuous
the Cauca and Magdalenavalleys, and on six months earlier (65% of marked frogs)
the Amazonian slopes of the Cordillera but were absent in January. Calling males
Oriental. Cochranand Goin(1970)reported were heard at night in pastures (calling on
a specimenfrom the isolated SierraNevada grass or low herbs) especially on rainy
de Santa Marta (Museum of Comparative nights. The call is a series of soft chuckles
Zoology4074). That specimenis a paratype "brarck, brarck, brarck" (6-8 notes per call
of E. insignitus Ruthven and the only ex- group). No amplectant pairs were seen.
ampleof E. w-nigrumknownfromthe Santa The distinct association with non-forested
Martas. I suspect a confusionof specimens habitats remained unchanged although
September 1979] HERPETOLOGICA 233

more frogs were seen up on vegetation Acknowledgments.-Support for fieldwork in-

(possibly a function of breeding activity). cluded grants from the Watkins Fund of the
Museum of Natural History, The University of
Discussion.-On the basis of similarities Kansas (1968), The Penrose Fund of the American
in morphologies, E. actites, E. lymani, and Philosophical Society (1970), and the University
E. w-nigrum appear to be sister species. In of Nebraska Research Council (1970, 1977). D.
view of their apparent closeness of relation- Cannatella, P. Fritts, T. Fritts, R. Henderson, and
M. Lynch aided me in fieldwork. Logistic support
ship, it is not surprising that they geograph- from F. Ortiz and E. del Piuo measurably aided
ically replace one another. In view of the fieldwork. For loan of specimens and/or provision
closeness of relationship and geographic of working space I thank W. Duellman, A. Grandi-
replacement, it would be expected that the son, W. Heyer, R. Inger, K. Klemmer, A. Kluge,
frogs would be similar ecologically. Data A. Leviton, H. Marx, C. Myers, R. Nussbaum, E.
Williams, G. Zug, and R. Zweifel.
for E. lymani are limited but extensive data
are available for E. actites and E. w-nigrum.
LITERATURECITED
Adult and juvenile E. w-nigrum are active
at night in contrast to the diurnal juveniles COCHRAN,D. M., AND C. J. GOIN. 1970. Frogs
and nocturnal adults of E. actites. E. w- of Colombia. U.S. Na'd. Mus. Bull. no. 288, 655
nigrum is closely associated with densely PP.
LYNCH,J. D. 1976. The species groups of the
wooded streams, whereas E. actites is a frog
South American frogs of the genus Eleuthero-
of open, watered microhabitats. The two dactylus (Leptodactylidae). Occas. Pap. Mus.
frogs also differ in calls-that of E. w- Nat. Hist. Univ. Kans. no. 61, 24 pp.
nigrum is loud and sounds like the call of SAVAGE,J. M. 1974. On the leptodactylid frog
many Ecuadorian Gastrotheca ("braaack, called Eleutherodactylus palmatus (Boulenger)
bonk, bonk, bonk, bonk") according to W. and the status of Hylodes fitzingeri 0. Schmidt.
Duellman. The two species may have over- Herpetologica 30:289-299.
lapping breeding seasons on the Pacific 1975. Systematicsand distributionof the
Mexican and Central American stream frogs
slopes of Ecuador but the available data are related to Eleutherodactylutsrugulosus. Copeia
limited (E. w-nigrum breeds from January 1975:254-306.
to April; E. actites has only been observed
to breed in January). Accepted: 27 November 1978
Thus E. actites can be distinguished from
its relatives by morphology, mating call, and Schoolof Life Sciences,The Universityof
ecology. Nebraska,Lincoln, Nebraska68588, USA