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OCCASIONAL PAPERS 
of the 0CT b Wb 
MUSEUM OF NATURAL HISTORY 
The University of Kansa^ NlvERSI 
Lawrence, Kansas 
NUMBER 38, PAGES 1-46 SEPTEMBER 10, 1975 
A REVIEW OF THE 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 
OF SOUTH AMERICA (LEPTODACTYLIDAE) 
Bv 
John D. Lynch 1 
Introduction 
Most eleutherodactyline frogs are relatively small and non- 
descript ( at least in preservative ) . However, several species found 
in forested habitats in northwestern South America and Central 
America are distinctive in having large, broad heads (head width 
45-63 percent of snout-vent length ) , prominent cranial crests, and in 
being comparatively large frogs (adult females 30-100 mm SVL). 
The superficial resemblance of these frogs to Ceratophrys has been 
cited (Dunn, 1944; Rivero, 1961). Most other eleutherodactyline 
frogs have "narrow" heads (HW/SVL = 30-43%), lack cranial crests, 
and do not exceed snout-vent lengths of 50 mm. Certain species 
match the character states of the broad-headed eleutherodactylines 
for one characteristic but not for all. For example, Eleutlierodac- 
tylus anomalus is more broad-headed than most species (HW/SVL 
= 41.8-48.3), large (adult females are larger than 50 mm SVL), but 
lacks cranial crests; E. curtipes and E. galdii have prominent cranial 
crests but are smaller frogs (adult females less than 40 mm SVL) 
with "narrow" heads; in some populations, E. fttzingeri is a large 
frog (adult females 60-75 mm SVL), but all populations have nar- 
row heads and lack cranial crests. 
The broad-headed eleutherodactylines include the type species 
of the genera Amblyphrynus Cochran and Goin, Ctenocranius Melin, 
1 Associate Professor of Zoology, School of Life Sciences, University of Nebraska, Lincoln, 
Nebraska 68508; Research Associate in Herpetology, Museum of Natural History, The 
University of Kansas, Lawrence, Kansas 66045. 
2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Limnophys Jimenez de la Espada, and Strabomantis Peters. No 
author has united all the species into a single genus although the 
idea seems to have been considered at various times (see Rivero, 
1961). Fourteen trivial names are available for broad-headed eleu- 
therodactylines. These are disposed in two genera: Amblyphrynus 
(ingeri Cochran and Goin) and Eleutherodactylus (biporcatus 
Peters, bufoniformis Boulenger, cornutus Jimenez de la Espada, 
florulentus Cope, gulosus Cope, koki Melin, macrocephalus Peracca, 
maussi Boettger, megacephalus Cope, napaeus Jimenez de la Espada, 
pelviculus Cope, rugosus Peters, and sulcatus Cope). 
In Central America, the broad-headed frogs occur from Hon- 
duras to Panama. Either two species (biporcatus and bufoniformis) 
are recognized or biporcatus is fragmented into three species — 
biporcatus (with a single putative synonym, napaeus), florulentus, 
and rugosus (with three putative synonyms, gulosus, megacephalus, 
and pelviculus). The Central American frogs are under study by 
J. M. Savage and further comment on them is restricted to South 
American populations. I consider the Central American broad- 
headed frogs as constituting two species, biporcatus and bufoni- 
formis. 
All names applied to South American specimens of broad-headed 
frogs are currently recognized, with the exception of napaeus. I do 
not consider napaeus a synonym of biporcatus. 
Historical Resume 
Peters (1864) named Strabomantis biporcatus from Veragua 
(= western Panama). He noted the relationships of this frog to 
Hylocles (= Eleutherodactylus), in that both groups are partially 
characterized by having "adhesive" discs on the digits, but separated 
them because Strabomantis had a broad head and narrow interorbital 
region. The subsequent history of the Central American taxa is sum- 
marized by Dunn (1931), Noble (1918), and Taylor (1952). I 
follow Dunn in recognizing two Central American species of "broad- 
headed" eleutherodactylines; biporcatus, florulentus, gulosus, mega- 
cephalus, pelviculus, and rugosus are considered conspecific. 
Jimenez de la Espada (1870) named Limnophys cornutus and 
L. napaeus from the Rio Suno near San Jose de Moti, Napo Province, 
Ecuador. He later (1872) considered Limnophys a synonym of 
Strabomantis, L. napaeus a synonym of S. biporcatus, and cornutus 
to be a valid species of Strabomantis. Both species described by 
Jimenez de la Espada had smooth skin on the venter and coarsely 
tuberculate skin on the dorsum. The holotype of L. napaeus was 
50 mm SVL; comparable sized biporcatus have nearly smooth skin 
above and I thus reject Jimenez de la Espada's synonymy of napaeus 
with biporcatus. 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 3 
Cope (1874) named Hylodes sulcatus from Nauta, Departa- 
mento Loreto, Peru, and considered it "evidently allied to Strabo- 
mantis biporcatus Peters." H. sulcatus differs most conspicuously 
from biporcatus, cornutus, and napaeus, in having coarsely areolate 
skin on the venter. 
Boulenger ( 1882) recognized three species of broad-headed eleu- 
therodactylines, biporcatus (including napaeus), cornutus, and 
sulcatus, but placed all of them in Hylodes (= Eleutherodactylus) . 
Boulenger did not have specimens of any of the named species avail- 
able; he had a single specimen (BMNH 69.7.25.11) from Bogota, 
Colombia, which he referred to cornutus and which served as partial 
basis for his description of the species. He presumed his specimen 
to be a young specimen of cornutus and thus recorded a mosaic of 
character states based on Jimenez de la Espada's account and 
BMNH 69.7.25.11. While recording Jimenez de la Espada's char- 
acteristic of broad, arched prevomerine dentigerous processes behind 
the choanae (not the character state in BMNH 69.7.25.11), he also 
recorded the skin of the venter as areolate. Boulenger also recorded 
the skin of the venter as granular in biporcatus, thus separating 
Cope's gulosus, megacephalus, and pelvicidus ( all broad-headed taxa 
from Central America ) from biporcatus. Peters' original description 
of biporcatus reads, "Eben so zeigt die Bauchseite kleine Warzchen, 
wahrend die Kehle glatt ist." and suggests a character state unlike 
that seen in sulcatus or in the BMNH specimen Boulenger referred 
to cornutus, both of which have coarsely areolate skin on the venter. 
However, Boulenger's error was incorporated into the literature; 
Nieden (1923) recorded the character state of the skin of the venter 
in biporcatus, cornutus, and sulcatus as "Unterseite gekornelt" or 
"Bauch gekornelt." 
Boettger (1893) named Hylodes maussi from Puerto Cabello, 
Venezuela; the species has coarsely areolate skin on the venter and 
was considered allied to cornutus and sulcatus. 
Boulenger (1896) named Hylodes bufoniformis from Buenaven- 
tura, Colombia. He did not compare bufoniformis with other broad- 
headed eleutherodactylines and some authors have not considered 
the species allied to the broad-headed complex but have suggested 
a relationship with the frogs of the fitzingeri-longirostris complex 
(group I of Cochran and Goin, 1970; binotatus group of Lynch, 
1973). 
Peracca (1904) named Hylodes macrocephalus from Valle San- 
tiago, Ecuador. This species was another of the frogs with coarsely 
areolate skin on the venter. Peracca considered it allied to sulcatus 
and distinguished the two taxa on the basis of head shape and the 
degree of cranial crest development. 
Peracca (1914) and Dunn (1944) reported additional Colombian 
specimens of an areolate-bellied frog with elongate supercilliary 
4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
tubercles as cornutiis. Dunn's specimen was later made the holotype 
of Amblyphrynus ingeri (Cochran and Goin, 1961). 
Melin (1941) named Ctenocranius koki from Taracua, Brasil. 
He considered koki allied to cornutus and maussi, and included 
cornutus, but not maussi, in the genus Ctenocranius. C. koki has 
areolate skin on the venter. 
Rivero (1961) regarded cornutus (sensu Boulenger) and maussi 
conspecific. He suggested that sulcatus and koki might prove to be 
conspecific with cornutus but noted that sulcatus was ". . . described 
as having an areolate belly." It is unclear if Rivero thought any of 
the others had smooth bellies; he recorded a granular venter in 
niaussi. 
My interest in these frogs began with the discovery that E. cornu- 
tus cornutus of Rivero was very much unlike the illustrations and 
descriptions given by Jimenez de la Espada (1870, 1875) but in 
agreement with the descriptions of Cope ( 1874), Melin ( 1941), and 
Peracca (1904). My interest was increased in 1968 when W. E. 
Duellman secured living specimens matching Jimenez de la Espada's 
description and figures of Limnophys cornutus. Additional speci- 
mens of this smooth-bellied frog with a elongate eyelid tubercle 
were found in the GOV and JAP collections at the National Museum 
of Natural History and in the collections at the American Museum of 
Natural History. The areolate-bellied frogs from Andean Colombia 
are juveniles with well-developed cranial crests and lacking digital 
pads. In most features they are identical to Amblyphrynus ingeri; 
the differences are assumed to be due to immaturity. The specimen 
reported by Cochran and Goin (1970) as the second specimen of 
Amblyphrynus ingeri has digital pads, a smooth venter, and numer- 
ous other differences from the description of Amblyphrynus ingeri; 
it is an adult female Eleutherodactylus biporcatus. 
No species of broad-headed eleutherodactylines, other than this 
latter specimen of E. biporcatus were previously known from the 
Pacific versant and lowlands of South America. Specimens of three 
species now have been found in the material at the American Mu- 
seum of Natural History, British Museum (Natural History), Na- 
tional Museum of Natural History, and the University of Kansas 
Museum of Natural History. Two of these species are smooth- 
bellied forms (E. cerastes new species and E. necerus new species) 
whereas the third is an areolate-bellied frog lacking digital pads 
(Amblyphrynus helonotus new species). 
Acknowledgments 
Specimens were loaned by the following curators and their re- 
spective institutions. The abbreviations following the name of the 
institutions are used throughout the text to identify particular speci- 
mens. James Bolhke, Academy of Natural Sciences, Philadelphia 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 5 
(ANSP); James R. Dixon, Texas Cooperative Wildlife Collection, 
Texas A and M University (TCWC); William E. Duellman, Univer- 
sity of Kansas Museum of Natural History ( KU ) ; Alice G. C. Grandi- 
son, British Museum (BMNH); Birgitta Hansson, Goteborgs Natur- 
historiska Museum (GNM); Rogert F. Inger and H. Marx, Field 
Museum of Natural History (FMNH); Richard Mount and Terry 
Schwaner, Auburn University Museum (AUM); Umberto Parenti, 
Museo de Istituto di Zoologia Sistematica della Universita di 
Torino (MZS); the late James A. Peters and George Zug, United 
States National Museum (USNM); Hobart M. Smith, University of 
Illinois Museum of Natural History (UIMNH); Charles F. Walker, 
University of Michigan Museum of Zoology (UMMZ); Ernest E. 
Williams, Museum of Comparative Zoology (MCZ); John Wright, 
Los Angeles County Museum ( LACM ) ; and Richard G. Zweifel and 
Charles W. Myers, American Museum of Natural History ( AMNH). 
William E. Duellman took photographs of living frogs, checked the 
Madrid collection for Jimenez de la Espada's types, and has gen- 
erously undertaken field work in my behalf in quest of these frogs. 
Charles W. Myers loaned a photograph of a rare species. The Uni- 
versity of Nebraska Research Council provided funds enabling me 
to study at the United States Museum and the British Museum 
(Natural History). James A. Peters and Alice Grandison provided 
working space at their respective institutions. Juan Leon provided 
a Spanish summary for the paper. My thanks go to all. 
Taxonomic Characteristics 
The primary characteristics of the broad-headed eleutherodac- 
tylines are head width and the presence of cranial crests. This com- 
bination of characteristics occurs in the nine species here termed 
"broad-headed eleutherodactylines" as well as in another telmato- 
biine genus, Proceratophrys ( Odontophrynini ) . The leptodactylids 
of the subfamily Ceratophryinae ( Lynch, 1971 ) have broad heads 
(HW/SVL = 45-65%) but lack cranial crests. 
Head width/ snout vent length (HW/SVL).— The range of this 
feature among eleutherodactyline frogs is 29.9-62.8%. The distribu- 
tion of HW/SVL values for eleutherodactyline frogs is bimodal; a 
pronounced mode is centered at about 36-38%, a smaller one at about 
49-50%. Percentages at around 50% apply to the frogs here termed 
"broad-headed"; the species and the ranges of HW/SVL for each are 
(N for each sample is given parenthetically following the range): 
A. helonotus new species .. _ 49.5-51.0(2) 
A. ingeri . 55.1-62.8(3) 
E. biporcatus _ . 47.9-54.3(27) 
E. bufoniformis _ . 44.4-58.2(22) 
E. cornutus 48.8-56.4(7) 
6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
E. cerastes new species 45.8-52.8(15) 
E. maussi . __ 44.3-54.9(10) 
E. necerus new species 46.2-49.7(3) 
E. sulcatus . .-- 45.6-53.3(15) 
The following three species of Eleutherodactylus are annectant be- 
tween "broad-headed" and "narrow." 
E. anomalus .. - 41.8-48.3(21) 
E. lymani ..... _ 35.7-44.5(16) 
E. taurus _ 40.8-45.3(8) 
The category of "narrow" head widths includes the following 
examples. 
E. achatinus _ ___. 33.5-43.2 ( 156) 
E. chloronotus .. 33.9-42.2(28) 
E. conspicillatus . 36.2-42.0(36) 
E. curtipes . -~~ 29.9-40.6(461) 
E. surdus 33.6-40.8(28) 
E. unistrigatus _ - 34.8-39.4(40) 
E. variabilis .. 30.7-37.2(20) 
E. w-nigrum - - 33.7-40.2(77) 
Phrynopus (13 spp) . 31.1-40.8(282) 
Syrrhophus (14 sp) . . 30.8-42.4(338) 
All of the "broad-headed" taxa have cranial crests, although those of 
E. bufoniformis are not as well-developed as those of some broad- 
heads, e.g., E. biporcatus or E. sulcatus. None of the annectant taxa 
have cranial crests whereas the occurrence of cranial crests among 
the frogs with "narrow" heads is sporadic — the presence of crests is 
interspecifically variable (e.g., in Phrynopus flavomaculatus, P. park- 
ed, and P. pereger). 
The South American specimens of broad-headed eleutherodac- 
ty lines appear to represent nine species. Each can be diagnosed on 
the basis of color patterns, proportions, and a suite of other charac- 
teristics. The color patterns are distinctive but difficult to use for 
grouping taxa. Most of the other characteristics can be used to 
group species and thus provide data for an analysis of intragroup 
relationships among broad-headed eleutherodactylines. 
In addition to coloration, the characteristics useful in species 
discrimination are as follows: 
1. — Texture of skin of dorsum. The arrangement of tubercles 
and folds, while difficult to describe, can be used in species 
recognition; the degree of tubercularity is ontogenetically 
variable within E. biporcatus. 
2. — Texture of skin of venter. The venter is either coarsely 
areolate (granular) or smooth. The character shows no 
intraspecific variation. 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 7 
3. — Eyelid tubcrculation. The eyelid is tubercular in all species 
but some forms have one or two elongate posterolateral eye- 
lid tubercles. 
4. — Tarsal fold. An inner tarsal fold is consistently present in 
some species. 
5. — Digital discs. Three character states are consistent. No 
digit bears discs; discs (on the ventral surface of the digit, 
bounded by a circumferential groove) are present on the 
toes but not the fingers; or, discs are present on the fingers 
and toes. 
6. — Ulnar tubercles. The posterolateral edge of the forearm may 
bear a series of large, crest-like warts or a series of smaller, 
less prominent tubercles. This feature varies with the size 
of the animal and is thus of limited value. 
7. — Supernumerary plantar tubercles. Prominent supernumer- 
ary plantar tubercles occur in the adults of only two species. 
Juveniles of at least one another species are known to have 
these tubercles but they are absent in adults. The feature 
is thus of limited value. 
8. — Lateral fringes on digits. If present, lateral fringes are most 
prominent on the toes. The lateral fringes are never flap- 
like, as in Hylodes (Elosiinae), but are easily recognized in 
juveniles and adults. 
9. — Basal webbing. This trait is recorded as "toes free" if the 
webbing does not enclose the basal subarticular tubercles 
of the toes, and "webbed" if the basal subarticular tubercle 
is enclosed. A single species has basal webbing. 
10. — Metatarsal tubercles. All specimens have two, and the inner 
is more prominent than the outer. Some species have an 
obscure outer tubercle and a prominent, laterally com- 
pressed inner tubercle, whereas others have a small, but 
prominent outer and a larger, non-compressed inner meta- 
tarsal tubercle. 
11. — Flaring of the lips. Only specimens from Venezuela and 
the Amazon basin have prominently flared lips as juveniles 
and adults. Adults and juveniles of all other taxa have little 
or no flaring of the lips. 
12. — Snout profile. In lateral profile the snout is either strongly 
sloping, weakly sloping, or truncate. The difference between 
"weakly sloping" and "truncate" is not consistent within a 
species. 
13. — Prevomerine processes. Two character states are evident — 
triangular in outline or broad and arched (lateral edge 
reaches to outer edge of choana). Ontogenetic variation 
seen is in one species: triangular as juveniles, arched as 
adults. The characteristic is thus of limited value. 
8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
14. — Vocal slits. The males of most species have vocal slits but in 
two species males evidently lack vocal slits. In each case, 
the males are small. Males are not available for all species 
and the characteristic is thus of limited value. 
15. — Cranial crests. All species have crests on the frontoparietal 
bones. In adults, the crests are lateral ( and largely parallel ) 
or parasagittal (and converging). In two species, there is 
some evidence of the crests extending onto the nasals. In 
most species, the crests are absent or obsolete in juveniles; 
the characteristic is thus of limited value. 
16. — Temporal region. The infratemporal fenestrum is occluded 
by bone ( squamoso-maxillary contact) in the holotype of 
Amblyphrynus ingeri. In all other specimens, including 
juvenile A. ingeri, the squamosal is not in contact with the 
maxillary. 
17. — Osteoderms. The holotype of A. ingeri has osteoderms in 
the skin of the dorsal surfaces. No other specimen, includ- 
ing juvenile A. ingeri, has detectable osteoderms. 
18. — Tympanum size. No difference in ear size among juveniles 
and males and females occurs in some species, whereas in 
other species (e.g., E. biporcatus) males have much larger 
tympana than do females. Because male specimens are 
lacking for some species and ontogenetic series are not avail- 
able for all species, the feature is of limited value. 
19. — Shank length. The variation of shank length within species 
renders this feature of questionable value. Some species 
have short legs (Amblyphrynus), whereas others are com- 
paratively long-legged (E. cornutus); but, the differences 
are not diagnostic. 
Characteristics in which a character state is unique to one species 
are of limited value in discerning relationships. For a characteristic 
to be of value, each character-state must be represented in two or 
more taxa. Likewise, characteristics in which the character-states 
change ontogenetically within a putative species are of little value 
unless comparisons are made only among individuals of the same 
sex and age. Therefore, of the above characteristics, only numbers 
2, 3, 4, 5, 8, 10, 11, and 12 are available for comparison of taxa. 
Texture of venter (2). — Cope (1866) distinguished Hylodes and 
Lithodytes on the basis of the texture of the skin of the venter. 
Under Cope's system, Hylodes included the frogs with digital discs 
having areolate skin on the venter and Lithodytes included those 
with discs and smooth skin on the venter. Jimenez de la Espada 
(1870) diagnosed Limnophys, and later (1872) St rabo mantis, on 
the basis of smooth skin on the venter. 
The broad-headed taxa with smooth skin on the venter are 
biporcatus, bufonijormis, cornutus, florulentus, gulosus, megacepha- 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 9 
lus, napaeus, pelviculus, and rugosus. Those with areolate skin on 
the venter are ingeri, koki, macrocephalus, maussi, and sulcatus. 
One undescribed species (helonotus) has areolate skin on the venter 
and two (cerastes and necerus) have smooth skin on the venter. 
Poorly preserved specimens of areolate-venter taxa sometimes 
superficially appear to have smooth skin on the venter. Examina- 
tion under magnification reveals the true character-state. Of greater 
difficulty is distinguishing a superficial areolation in smooth-venter 
taxa. In these examples the "warts" appear to be due to an effect of 
preservatives and are spinule-like rather than the pavement-like 
texture seen in frogs having areolate skin on the venter. The prob- 
lem is most severe in small frogs ( less than 25 mm SVL ) . 
Eyelid tubercles (3). — All species of broad-headed eleutherodac- 
tylines have tubercles on the upper eyelids. The character-states 
used in distinguishing taxa reflect not the presence of tubercles but 
the relative lengths of the tubercles. All specimens here assigned to 
cerastes, cornutus, and ingeri have elongated tubercles (Fig. 1) at 
Fig. 1. — Lateral profiles of heads of EleutJierodactylus cornutus (left, KU 
123448) and E. maussi (right, UMMZ 113940, AB 4496). 
the posterolateral corner of the upper eyelids. The tubercles are 
evident in juveniles as well as large adults. Jimenez de la Espada 
(1870) recorded shorter tubercles in napaeus than in cornutus; he 
illustrated (1875) only cornutus, so it is not possible to determine if 
napaeus had distinct horn-like eyelid tubercles such as occur in 
cerastes, cornutus, and ingeri or whether the eyelid tuberculation 
was wart-like, such as that seen in biporcatus, bufoniformis, koki, 
macrocephalus, maussi, or sulcatus (Fig. 1). 
Tarsal fold (4). — An inner tarsal fold is found in the Amazonian 
(cornutus, koki, inacrocephalus, sulcatus) and Andean (ingeri, maus- 
si) taxa but not in those from the Pacific versant or lowlands or those 
from Central America. 
Digital discs (5). — Two species (helonotus and ingeri) lack discs 
on the digits. The disc is the structure on the ventral surface of the 
digital tip surrounded by a groove; the groove is most distinct at the 
tip of the digit. I use the term "digital pad" for any apical dilation 
of the digit. The nominal species koki, macrocephalus, and sulcatus 
have discs on the toes but none on the fingers although the fingers 
10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
may have apical swelling and some dilation (Fig. 2). All other 
broad-headed taxa have discs on the fingers and toes. In all species 
the tips of the toes are broader (and pad-like) than those of the 
fingers. 
Fig. 2. — Palmar views of hands of Eleutherodactylus biporcatus ( left, KU 
108426) and E. sulcatus (right, KU 126171). 
Supernumerary plantar tubercles (7). — koki, macrocephalus, 
maussi, and sulcatus have prominent supernumerary plantar tuber- 
cles; I also found less prominent tubercles on the plantar surfaces of 
young itigeri. No adult broad-headed eleutherodactyline except 
maussi and sulcatus has supernumerary plantar tubercles. 
Lateral fringes on digits (8). — The fingers lack basal webbing, 
and even when present, lateral fringes on the fingers are difficult to 
detect (Fig. 2). Lateral fringes on the toes are readily detected. 
Most South American broad-headed taxa have lateral fringes; fringes 
are absent in biporcatus and cerastes. 
Metatarsal tubercles (10). — All specimens examined have two 
metatarsal tubercles and the inner is larger than the outer. The inner 
metatarsal tubercle is strongly compressed laterally and the outer 
obscure in biporcatus, bufoniformis, and necerus. In the other 
species the inner tubercle is not compressed, or has only slight lateral 
compression, and the outer is more prominent (Fig. 3). 
Flaring of the lips (11). — The loreal region is more vertical in 
biporcatus, bufoniformis, cerastes, cornutus, ingeri, and necerus, 
than in koki, macrocephalus, maussi, and sulcatus. In the latter 
group of taxa, the snout is strongly sloping in lateral profile ( Fig. 1 ) , 
a feature not seen in the other broad-headed taxa. Associated with 
flaring of the lips is the flaring of the posterior maxillary arch. The 
frogs with posterior flaring have an obliquely oriented tympanum. 
The increase in slope of the temporal region increases ontogeneti- 
cally (Fig. 4) so that young frogs have nearly vertical tympana 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 
11 
Fig. 3. — Proximal portions of feet of (A) Eleutherodactylus cornutus, KU 
123448, and ( B ) E. biporcatus, KU 108425, and plantar level views of meta- 
tarsal tubercles of (C) E. cornutus, KU 123448, and (D) E. biporcatus, KU 
108425. Arrows point to outer metatarsal tubercles. 
whereas large adults (especially females) have markedly oblique 
tympana. The slope of the tympanum was used by Peracca ( 1904 ) 
to distinguish marcocephalus from sulcatus; the distinction is size- 
related — the holotype of macrocephalus is 35 mm SVL, that of 
sulcatus 47 mm SVL. 
Prevomerine processes (13). — All specimens of broad-headed 
eleutherodactylines have prevomerine teeth, but the shape of the 
prevomerine processes varies from triangular to arched ( Fig. 5 ) . The 
processes are triangular in all Central American frogs (nominal 
species biporcatus, fiorulentus, gulosus, megacephalus, pelviculus, 
and rugosus) as well as bufoniformis, ingeri, and the small specimens 
of necerus. The processes are arch-like (extending laterally to the 
middle or external edge of the choanae) in adult necerus, and all 
specimens of cornutus, helonotus, koki, macrocephalus, maussi, na- 
paeus, and sulcatus. The holotype of sulcatus and a second Peruvian 
specimen (AMNH 43391) are the only exceptions; in both speci- 
mens, the lateral extent of the prevomerine processes is the median 
12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Fig. 4. — Outline drawings of anterior view of heads of E. maussi, (A) 
AMNH 70549, 67.9 mm SVL; (B) AMNH 70546, 36.4 mm SVL; (C) AMNH 
70548, 25.0 mm SVL; (D) AMNH 70542, 22.0 mm SVL; and (E) AMNH 
70536, 12.7 mm SVL. 
border of the choanae. The narrow processes of these two examples 
are considered abnormal, but the occurrence of such variation miti- 
gates the value of the feature. 
Triangular prevomerine processes are found in most frogs of the 
binotatus group. No eleutherodactyline frogs from South America 
except some broad-headed taxa have arched processes. Arched 
processes are known among West Indian Eleutherodactylus ( inopta- 
tns and ricordii groups ) and in most Leptodactylus. 
Cranial crests (15). — The cranial crests are borne on the fronto- 
Fig. 5. — Anterior palates of Eleutherodactifhis bufoniformis (left, KU 
113787) and E. sulcatus (right, AMNH 43136). 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 
13 
Fig. 6. — Roofing bones (and portions of neurocrania) with cranial crests 
of (A) Eleutherodactylus bufoniformis, KU 80621, (B) Amblyphrynus helo- 
notus, BM 1970.178, (C) E. biporcatus, LACM 73157, (D) E. cerastes, KU 
144992, (E) E. sulcatus, AMNH 52856, and (F) A. ingeri, AMNH 39979. 
parietals and are either lateral (Figs. 6C, E and F) or parasagittal 
(Figs. 6A, B, and D). Most species have lateral crests; parasagittal 
crests occur in bufoniformis, cerastes, helonotus, and necerus. In 
young biporcatus and sulcatus the crests are somewhat parasagittal; 
in larger frogs they become lateral. Crests are poorly developed and 
nearly indistinguishable in juveniles and young males of maussi; in 
general, crests of males are less well-developed than those of females. 
In ingeri and, to a lesser extent, sulcatus, the crests continue onto 
the nasal bones. I did not find crests on the nasals of any other 
specimens examined, but Cope (1875) reported the crests extending 
onto the nasals in the holotype of Lithodytes gulosus. Posteriorly, 
the crests terminate in an enlarged knob or boss in large specimens. 
The boss is most prominent in large biporcatus (Fig. 6C) and in 
juvenile ingeri ( Fig. 6F ) . In all examples with crests, the crests are 
higher posteriorly and approximate an elevated boss. 
The otic ramus of the squamosal forms a crest in some examples. 
14 
OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Fig. 7. — Left: reconstruction of skull of Arriblyphrynus ingeri based on 
radiographs (FMNH 81915). Top right: outline drawing of head of Eleu- 
therodactylus biporcatus ( hatching indicates position and shape of cranial 
crests). Lower right: lateral view of head of E. biporcatus ( LACM 73157) 
showing extent of bones and position of tympanum. 
The otic crest is most pronounced in Amblyphrynus ingeri (Fig. 7) 
but detectable crests are also found in larger examples of maussi and 
sulcatus. A crest-like keel is found on the otic ramus of the squa- 
mosal in large adult female biporcatus; the keel is not parallel to 
the cranial crests as in ingeri ( Figs. 6-7 ) and sulcatus. 
Key to Broad-headed Eleutherodactylines 
1. Digits lacking discs 2 
At least toes bearing discs (defined by circumferential grooves) 3 
2. Tarsus bearing fold along inner edge; cranial crests prominent 
ending posteriorly in large boss Amblyphrynus ingeri 
Tarsus lacking folds; cranial crests obscure, no enlarged boss pos- 
teromedial to eyes Amblyphrynus helonotus 
3. Skin of venter coarsely areolate 4 
Skin of venter smooth 5 
4. Fingers lacking discs; posterior surface of thighs with large pale 
spots Eleutherodactylus sulcatus 
Fingers bearing discs; posterior surface of thighs uniform brown 
Eleutherodactylus maussi 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 15 
5. Upper eyelid bearing elongate tubercle at posterolateral corner 6 
Upper eyelid lacking elongate tubercles 7 
6. Tarsal fold present; toes bearing lateral fringes 
Eleiitherodactylus cornutus 
Tarsus lacking folds; toes lacking lateral fringes . 
Eletitherodactylus cerastes 
7. Toes lacking lateral fringes Eleiitherodactylus biporcatus 
Toes bearing lateral fringes 8 
8. Basal subarticular tubercles of toes enclosed by webbing 
Eleiitherodactylus bufoniformis 
Basal subarticular tubercles of toes not enclosed in toe webbing .. 
Eleiitherodactylus necerus 
Systematic Accounts 
The published descriptions of E. biporcatus, E. bufoniformis, E. 
maussi, and E. sulcatus are adequate and not duplicated here. For 
these taxa I have provided an extensive diagnosis and definition as 
well as references to published descriptions. Because altitudinal 
data are not available for the majority of specimens, I have used 
general statements for the elevational distributions, viz., low (0-800 
m), moderate (800-1600 m), and intermediate (1600-2500 m). 
The fourteen nominate species of broad-headed eleutherodac- 
tyline frogs and three undescribed species are placed in two genera 
— Amblyphrynus and Eleiitherodactylus. 
Amblyphrynus Cochran and Goin 
Diagnosis. — Eleutherodactyline frogs with simple digits (no pads or discs; 
terminal phalanges knobbed) and broad-heads (HW/SVL: 50-63%). 
These frogs have areolate skin on the venter, the first finger longer than the 
second, lateral fringes on the digits, no webbing of the digits, prominent prevo- 
merine dentigerous processes, prominent, externally visible, tympana, cranial 
crests, and a cartilaginous omosternum. The nasals and frontoparietals are in 
broad median contact. 
Type-species. — A. ingeri Cochran and Goin, by original desig- 
nation. 
Content. — Two species found in northwestern South America. 
Amblyphrynus ingeri Cochran and Goin 
Fig. 8 
Hylodes cornutus (part): Boulenger, 1882:220; Peracca, 1914:107. 
Eletitherodactylus cornutus: Dunn, 1944-26; Cochran and Goin, 1970:442. 
Amblyphrynus ingeri Cochran and Goin, 1961:543; 1970:360. [Holotype. — 
FMNH 81915, 8 km S Gachala, San Isidro, Cundinamarca, Colombia, 2350 
m]. 
16 
OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Diagnosis. — A large eleutherodaetyline frog (one adult $ 50 mm SVL) 
with heavy lateral cranial crests on the frontoparietals; skull with extensive exo- 
stosis (nasals, frontoparietals, maxillae, squamosals); squamosal broadly con- 
tacting maxilla, no opening between squamosal, maxilla, and quadratojugal in 
adult female; prevomerine dentigerous processes triangular in outline, postero- 
medial to choanae; skin of dorsum bearing osteoderms; skin of dorsum weakly 
tuberculate in adult female (no large flat warts), coarsely tuberculate in juve- 
niles; upper eyelid bearing elongated tubercle on posterolateral surface; tarsal 
fold present; venter uniform brown. 
Description. — Cochran and Goin ( 1961, 1970) have described the 
holotype (the only known adult). A description of juvenile frogs 
(20.0-25.0 mm SVL) is provided here. 
Head broader than body, wider than long; head width 58.9-62.8% 
(x = 60.8) SVL; snout round in dorsal view, short and truncate in 
lateral profile ( Fig. 8 ) ; length of eye greater than distance between 
eye and nostril; canthus rostralis sharp, straight or slightly concave; 
loreal region somewhat concave, sloping gradually to lips; lips flared; 
nostrils protuberant, directed laterally; tip of snout not bearing 
proboscis or ridge, not extending much beyond lower jaw; inter- 
orbital region concave, edges of frontoparietals enlarged to form 
cranial crests; interorbital region broader than width of upper eye- 
lid; upper eyelid 75.0-88.9% (x = 80.3) interorbital distance; upper 
* ■ ■*; . . t 
^> J 
,? n 
I- 
-^v*-- 
Fig. 8. — Juvenile Amblyphrynus ingeri, AMNH 39979. 
eyelid bearing elongate superciliary tubercles; cranial crests termi- 
nate posteriorly in bosses lying posteromedially to eyes; anteriorly 
they form canthus rostralis; tympanum prominent, exposed, slightly 
higher than long, its horizontal diameter 56.1-66.7% length of eye in 
females, 84.3% in one male; supratympanic fold present, bearing 
tubercles; edges of lower jaw tuberculate; tongue large, fleshy, pos- 
terior one-fifth free, not notched posteriorly; choanae relatively small, 
oval, completely visible when roof of mouth is viewed from directly 
below; prevomerine dentigerous processes present, round, each 
BROAD-HEADED ELEUTHERODACTYLIXE FROGS 17 
bearing a clump of 4-5 teeth, each process about twice as large as a 
choana; males lacking vocal sac and vocal slit. 
Skin of dorsum pustular and bearing enlarged ridges and tu- 
bercles; tubercles prominent on eyelids, on interorbital region, and 
tending to form ridges on back; limbs tuberculate and ridged; tuber- 
cles and ridges less prominent on upper arm and thighs than on 
forearm, shank and tarsus; skin of venter coarsely areolate; discoidal 
folds indistinct, terminating posteriorly well anterior of thighs; shank 
52.0-54.5% (x = 53.4) SVL; outer edge of forearm bearing enlarged 
ulnar tubercles which tend to coalesce to form ulnar ridge; no 
enlarged tubercle on elbow; two palmar tubercles, outer and median 
largely fused (Fig. 9); thenar tubercles as large as subarticular 
Fig. 9. — Palmar view of hand of Amblyphrynus ingeri (AMNH 39978). 
Line equals 1 mm. The fourth finger is enlarged 2x at the left. 
tubercles, pale in color; subarticular tubercles round, simple, some- 
what conical; no supernumerary tubercles on digits; digits bearing 
slight lateral fringes; tips of digits not enlarged to form distinct pads, 
lacking circumferential groove at tips ( Fig. 9 ) ; first finger distinctly 
longer than second, heel not bearing enlarged tubercles; outer edge 
of tarsus bearing a row of enlarged tubercles extending from heel to 
outer metatarsal tubercle; inner surface of tarsus bearing a weak 
inner tarsal fold extending from inner metatarsal tubercle proximally 
for about % to % length of tarsus where it is replaced by a row of 
enlarged tubercles; inner metatarsal tubercle not compressed, about 
three times as long as wide and about three times as large as round, 
non-conical outer metatarsal tubercle; plantar surface bearing nu- 
merous supernumerary tubercles arranged in rows approximately 
corresponding to metatarsal bones; subarticular tubercles of toes 
smaller than those of fingers, round, flat, simple; tips of toes bearing 
small pads, about as long as wide; toes bearing ill-defined lateral 
IS 
OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
fringes but lacking webbing; outer edge of foot and fifth toe bearing 
a tuberculate ridge. 
Coloration in preservative. — Body brown with dark brown to 
black markings (canthal stripe, supratympanic stripe, labial bars, 
bars on limbs and edges of scapular ridges); posterior surface of 
thigh pale brown with numerous small, pigmentless spots; anterior 
face of thigh and lower flanks bear small pigmentless spots. 
Distribution. — Andes of Colombia at intermediate elevations 
(Fig. 10). Two literature records of Eleutheroclactyhis cornutus 
based on specimens not seen by me are believed to apply to A. 
ingeri. These are Peracca's (1914) frog from Camelia, Depto. An- 
tioquia Colombia, 1720 m, and Cochran and Goin's ( 1970 ) frog 
A A. ingeri 
▼ E.biporcatus 
• E. cerastes 
■ E.maussi 
• E.sulcatus 
400 KMS 
! 
68 
i_ 
64 
i 
8 
Fig. 10. — Map of northwestern South America showing localities at which 
Amblyphrynus ingeri, Eleuiherodactylus biporcatus, E. cerastes, E. maussi, and 
E. sulcatus have been found. Open triangle represents an unverified locality 
for A. ingeri. 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 19 
from Valdivia, Depto. Antioquia, Colombia (based on FMNH 69742, 
apparently lost). 
Remarks. — In comparison to the adult female, the juveniles are 
more coarsely tuberculate and have less flaring of the lips. The 
terminal phalanges of the juvenile frogs bear more pronounced 
lateral processes than do those of the holotype. The juveniles do not 
have detectable osteoderms and do not have the lower temporal 
region occluded by bone. In spite of these differences, which I pre- 
sume to reflect immaturity, the smaller, juvenile, specimens from 
Andean Colombia are considered conspecific with the adult female 
holotype of A. ingeri. 
Cochran and Goin ( 1970) referred an adult female eleutherodac- 
tyline (FMNH 54591) from the Pacific versant of Colombia to A. 
ingeri and noted certain differences between this specimen and the 
holotype. 
FMNH 54591 is a moderate-sized female (78 mm SVL) of E. 
biporcatus (females 65-110 mm SVL) and certainly not conspecific 
with the holotype of Amhlyphrynus ingeri. A. ingeri has eight 
procoelous presacral vertebrae, lacks a vertebral shield, has widely 
separated cervical cotyles, and a bicondylar sacro-coccygeal articu- 
lation. The transverse processes of the anterior presacral vertebrae 
are somewhat longer than those of the posterior presacral vertebrae 
and the non-dilated sacral diapophyses (which are deflected pos- 
teriorly). The ilia are of the leptodactyline type (Lynch, 1971) in 
having a dorsolateral ilial prominence (not spike-like) and an ilial 
crest. The pectoral girdle is arciferal, cartilaginous pre-zonal and 
post-zonal elements are present, and the clavicles and coracoids are 
neither slender nor massive and curved. 
The most distinctive feature of A. ingeri is the absence of an 
infratemporal fenestrum (Fig. 7). In other leptodactylids having 
maxillary-squamosal contact (Caudiverbera, Ceratophrys, Cijclorana 
australis, Lepidobatrachus, Megaelosia, and Proceratophrys) there 
is a prominent infratemporal fenestrum. The infratemporal fenestrae 
are small or absent in Hemiphractus (Hylidae), Genijophnjne (mi- 
crohylidae), and Ceratobatrachus (Ranidae), but I know of no 
other frogs that lack an infratemporal fenestrum. 
Amblyphrynus helonotus new species 
Fig. 11 
Holotype. — BMNH 1970.178, collected at the Rio Pitzara, Pichincha Prow, 
Ecuador, by M. Olalla. 
Paratype. — USNM 195784, collected at Mindo, Pichincha Prow, Ecuador, 
by M. Olalla, 24-31 May 1959. 
Diagnosis. — A large eleutherodactyline frog ( $ 9 60-70 mm SVL) with 
low, parasagittal cranial crests on the frontoparietals; skull with little exostosis; 
squamosal not contacting maxilla; prevomerine dentigerous processes arch-like, 
extending laterally to outer edge of choanae; dorsal skin lacking osteoderms; 
20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
skin of dorsum tuberculate bearing large flattened warts; upper eyelid lacking 
elongate tubercles; no tarsal fold; venter blotched with black or dark brown. 
Description. — Head as broad as or broader than body; head 
wider than long; head width 49.5-51.0% SVL; snout round in dorsal 
view (Fig. 11), truncate in lateral profile; snout moderately short, 
eye-nostril distance 93.8-96.1% eye length; canthus rostralis obtuse, 
not sharp, weakly concave; loreal region flat to weakly concave, 
sloping gradually to flared lips; interorbital space furrowed, low 
cranial crests present, not extending onto nasals and not ending in a 
prominent boss; upper eyelid width 143.4-167.4% IOD; supratym- 
panic fold present, indistinct; tympanum prominent, higher than 
long, its length 49.7-57.0% eye length, separated from eye by 1/2 to 
2 times width of tympanum; tongue about as long as wide, weakly 
notched posteriorly, posterior l A to % not adherent to floor of mouth; 
choanae small, not concealed by palatal shelf of maxillae; prevo- 
merine dentigerous processes present, arched, posterior to choanae; 
each process 3 to 4 times width of a choana; processes narrowly 
separated, extending to lateral edge of choana. 
Skin of dorsal surfaces tuberculate, tubercles heterogeneous 
(Fig. 11); a pair of indistinct canthal folds present; no enlarged 
"horn-like" tubercles on eyelids; a pair of )( shaped folds in scapular 
region; skin of venter coarsely areolate; no dorsolateral folds; dis- 
coidal folds not apparent; ulnar tubercles present; two palmar tuber- 
cles, outer largest, bifid; supernumerary palmar tubercles prominent, 
flattened; subarticular tubercles round, simple, non-conical; fingers 
Fig. 11. — Amblyphrymis helonotus (USNM 195784, paratype). 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 
21 
bearing weak lateral fringes; tips of fingers rounded, lacking pads; 
thumb longer than second finger. 
Heel and inner tarsus lacking tubercles or folds; outer edge of 
tarsus bearing row of small tubercles; two metatarsal tubercles, inner 
weakly compressed, twice as long as wide, 2 to 3 times size of 
rounded, non-conical outer metatarsal tubercle; no supernumerary 
plantar tubercles; toes bearing lateral fringes but lacking basal 
webbing; subarticular tubercles round, conical, simple, smaller than 
those of fingers; tips of toes not expanded, lacking pads. 
In preservative, A. helonotus is brown above with indefinite black 
or dark brown markings on the tubercles and folds. No distinct 
canthal or supratympanic stripes are evident. The face is barred 
with broad brown labial bars separated by thin cream lines. The 
venter is cream with dark brown blotches. The throat is dark brown 
with a few darker spots at the edge of the lip. The undersides of the 
limbs are dark brown with cream spots. The groin is dark brown 
and lacks a pattern. The limbs are barred with dark brown; the bars 
are about as wide as the brown interspaces. The bars on the thigh 
continue down the anterior face of the thigh and coalesce ventrally 
Fig. 12. — Map of Ecuador showing localities at which Amblyphrynus 
helonotus, Eleutherodactylus cornutus, and E. necerus have been found. Open 
circle represents a literature record for E. cornutus. 
22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
with the reticulation on the ventral surfaces of the thighs. The 
posterior surfaces of the thighs are brown with black reticulations. 
Measurements in mm. — The measurements of the type are listed 
first and the corresponding measurement of the paratype is listed in 
parentheses. SVL 69.6 (60.6), shank 31.0 (26.6), head width 35.5 
(30.0), head length 26.0 (23.4), eyelid width 7.6 (7.2), IOD 5.3 
(4.3), tympanum length 3.8 (3.6), eye length 7.6 (6.4), eye-nostril 
distance 7.4 (6.0). 
The holotype is a gravid female ( ovarian eggs yellow, 3-4 mm in 
diameter, heavily convoluted oviducts). The paratype is a young 
female (ovarian eggs white, %-l mm in diameter, weakly convoluted 
oviducts ) . 
Etymology. — Greek, helos -f- notos, wart -f- back; in reference 
to the large warts on the dorsum and flanks. 
Distribution. — Known only from the Pacific versant of Ecuador 
at low and moderate elevations (Fig. 12). 
Remarks. — A. helonotus differs markedly from A. ingeri. Com- 
parable intrageneric variation occurs within Cyclorana, Pyxicepha- 
lus, and Rhacophorus. I view the degree of relation between the 
two species of Amblyphrynus as roughly equivalent to that between 
Eleutherodactylus biporcatus and E. sulcatus, congeneric, but in 
different species groups. 
Eleutherodactylus Dumeril and Bibron 
Diagnosis. — Eleutherodactyline frogs with complex digits (discs present, 
supported by T-shaped terminal phalanges, at least on toes), teeth on maxillae, 
premaxillae, and prevomers, and lacking rows of tubercles on the plantar 
surfaces. 
Frogs of this genus are diverse. Most species have narrow heads 
(HW/ SVL: 30-42%), lack webbing of the toes, and lack a fronto- 
parietal fontanelle. The various species groups differ in head width 
(two groups with broad heads, biporcatus and sulcatus), texture of 
the venter (smooth vs areolate), finger length (first finger shorter 
than second vs longer than second), notching of the skin above the 
finger discs (notched in southeastern Brasilian Eleutherodactylus. 
and some partially studied osteological features (fusion of fronto- 
parietals and prootics, overlap of pterygoid and parasphenoid, size 
of prevomers, and shape of zygomatic ramus of squamosal). The 
tympana may be prominent, partially or wholly concealed, or 
absent. Cranial crests occur sporadically, mostly within the broad- 
headed groups and in a number of Andean species. 
Type-species. — Hylodes martinicensis Tschudi, by monotypy. 
Content. — Four hundred and thirty names have been proposed; 
generally 200 to 250 taxa are recognized. 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 
23 
Eleutherodactylus biporcatus (Peters) 
Strabomantis biporcatus Peters, 1864:405 [Type-locality — Veragua, Panama]. 
Hylodes rugosus Peters, 1874:610 [Type-locality — Chiriqui, Panama]. 
Lithodytes megacephalus Cope, 1875:110 [Type-locality — Pico Blanco, Costa 
Rica, 6000 ft.]. 
Lithodytes gulosus Cope, 1875:112 [Type-locality — Pico Blanco, Costa Rica, 
6000 ft.]. 
Lithodytes pelviculus Cope, 1877:89 [Type-locality — west coast of Central 
America]. 
Hylodes rugosus: Boulenger, 1882:205. 
Hylodes gulosus: Boulenger, 1882:211. 
Hylodes biporcatus: Boulenger, 1882:220. 
Hylodes megacephalus: Brocchi, 1882:57. 
Lithodytes florulentus Cope, 1893:336 [Type-locality — Boruca, Costa Rica]. 
Hylodes florulentus: Gunther, 1901:234. 
Eleutherodactylus rugosus: Noble, 1918:329. 
Eleutherodactylus biporcatus: Dunn, 1931:410. 
Eleutherodactylus florulentus: Taylor, 1952:765. 
Diagnosis. — A large Eleutherodactylus (adult 9 9 65-110 mm SVL) with a 
broad head ( HW/SVL = 47.9-54.3%); snout truncate in lateral profile; lips not 
flared; frontoparietals bear moderate lateral crests terminating posteriorly in 
large boss, normally not extending onto nasals; tympanum 80.0-89.8% eye length 
in males, 43.6-50.2% in females; prevomerine dentigerous processes triangular in 
outline, posteromedial to choanae; males with vocal sac and slits; ulnar tubercles 
Fig. 13. — Broad-headed Eleutherodactylus. (A) E. biporcatus, LACM 
73154, (B) E. bufoniformis, LACM 73248, (C) E. necerus, USNM 195799, 
(D) E. cerastes, USNM 195793, (E) E. sulcatus, AMNH 42293, and (F) 
E. maussi, AMNH 70557. 
24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
not or only weakly developed; two palmar tubercles, outer largest, bifid; fingers 
lacking lateral fringes; fingers bearing discs at tips, discs narrow, but broader 
than fingers; thumb longer than second finger; skin of dorsum moderately 
tuberculate in young, becoming smooth with scattered tubercles in large adults; 
upper eyelid tuberculate, no tubercle elongate; skin of venter smooth; tarsus 
lacking fold along inner margin; inner metatarsal tubercle strongly compressed 
laterally; outer metatarsal tubercle small and obsolete; plantar surface lacking 
supernumerary tubercles; toes lacking lateral fringes and basal webbing; toes 
bearing moderate-sized discs and pads, pads wider than long; posterior surface 
of thigh black or black with pale areas (posterior extensions of pale areas on 
dorsal surface of thigh ) ; venter usually reticulated with brown. 
Descriptions. — In addition to the original descriptions (Cope, 
1875, 1877, 1893, and Peters, 1864, 1874), descriptions are available 
in Noble (1918) and Taylor (1952, 1955), the latter including 
illustrations. 
Distribution. — Eastern Honduras and Nicaragua through low 
and moderate elevations to western Colombia (sea-level to about 
1000 m). 
Remarks. — Jimenez de la Espada (1872) incorrectly referred 
Limnophys napaeus to the synonymy of E. biporcatus. The holotype 
of napaeus is lost, but two features cited by Jimenez de la Espada 
suggest that napaeus is a synonym of cornutus and not of bipor- 
catus: the holotype of napaeus was 50 mm SVL, had coarsely tuber- 
culate skin of the dorsum and broad, and arched prevomerine 
dentigerous processes (Jimenez de la Espada, 1870). Comparable 
sized biporcatus have nearly smooth skin on the dorsum (Fig. 13) 
and triangular prevomerine processes. Accordingly, Limnophys na- 
paeus Jimenez de la Espada is removed from the synonymy of 
Eleutherodactylus biporcatus (Peters). 
The specimen (FMNH 54591) reported as Amblyphrynus ingeri 
by Cochran and Goin (1970) is a normal female E. biporcatus and 
in no way departs from the diagnosis given above. 
Eleutherodactylus bufoniformis (Boulenger) 
Hylodes bufoniformis Boulenger 1896:19 [Type-locality — Buenaventura, Valle, 
Colombia]. 
Eleutherodactylus bufoniformis: Dunn, 1931:410. 
Diagnosis. — A large Eleutherodactylus (adult females 52-94 mm SVL) with 
a broad head ( HW/SVL = 44.4-58.2%); snout truncate in lateral profile; fronto- 
parietals bear low parasagittal crests not extending onto nasals; tympanum 2/5 
to M eye length in females, M to % eye length in males; prevomerine dentigerous 
processes triangular in outline, posteromedial to choanae; males with vocal sac 
and slits; ulnar tubercles present, not enlarged; two palmar tubercles, outer 
largest, bifid; fingers lacking lateral fringes; digit tips bearing pads and discs, 
pads wider than finger; thumb longer than second finger, skin of dorsum tuber- 
culate and bearing short ridges in young as well as adult frogs; upper eyelid 
tuberculate, no tubercles elongate; skin of venter smooth; tarsus lacking inner 
fold ( or having a weak inner tarsal fold ) ; two metatarsal tubercles, inner largest, 
laterally compressed, outer obscure, small; no supernumerary plantar tubercles; 
toes bearing lateral fringes and basally webbed ( webbing encloses basal sub- 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 25 
articular tubercles ) ; toes with discs, pads broader than long, larger than those 
of fingers; posterior surfaces of tliigbs brown with small cream spots; dark color 
of lower flanks grades onto venter; venter mottled with brown. 
Descriptions. — Boulenger's ( 1896 ) brief original description is 
amplified by Cochran and Goin's ( 1970 ) . 
Distribution. — Low elevations, from extreme western Panama 
onto the Choco lowlands of Colombia. 
Eleutherodactylus cerastes new species 
Fig. 14 
Holotype. — USNM 195785, an immature female, collected at Palma Real, 
Pichincha Prov., Ecuador, collected June 1955 by M. Olalla. 
Paratypes.—KU 144992-93, El Tambo, La Costa, Depto. Cauca, Colombia, 
1000 m, KU 144994, El Tambo, Munchique, Depto. Cauca, Colombia, 1000 m; 
AMNH 88062, 13 km W Dagua, Rio Anchicaya drainage, Valle, Colombia, 850- 
1200 m; USNM 195786-88, Lita, Rio Mira, Imbabura Prov., Ecuador; USNM 
195789-93, Pachijal, Pichincha Prov., Ecuador; USNM 195794, Palma Real, 
Pichincha Prov., Ecuador. 
Referred specimens. — BMNH 1910.7.11.61-62, Pueblo Rico, Depto. Cauca, 
Colombia, 1580 m; USNM 195795, Lita, Rio Mira, Imbabura Prov., Ecuador; 
USNM 195796(3), Rio Pitzara, Pichincha Prov., Ecuador; USNM 195797, road 
from Pacto to Rio Guayllabamba, Pichincha Prov., Ecuador. 
Diagnosis. — A large Eleutherodactylus (2 adult 9 9 46.0-55.8 mm SVL); 
snout truncate or weakly sloping in lateral profile, lips weakly flared in large 
adults; frontoparietals bearing low lateral crests, not ending in bosses, not 
extending onto nasals; tympanum Ja to J« eye length in females, Y* to 3/5 eye 
length in males; prevomers with broad, arched processes, extending laterally to 
lateral edge of choana; males lacking vocal sac; ulnar tubercles present, not 
enlarged; two palmar tubercles, outer largest and bifid; fingers lacking lateral 
fringes; digit tips bearing narrow pads; thumb longer than second finger; skin 
tuberculate and ridged above in young and adult individuals; upper eyelid 
bearing elongate tubercle on posterolateral corner; skin of venter smooth; tarsus 
lacking folds, outer edge bearing row of small tubercles; two metatarsal tuber- 
cles, inner laterally compressed, outer small, rounded; no supernumerary plantar 
tubercles; toes lacking lateral fringes; toes with pads, broader than long, much 
larger than those of fingers; posterior surfaces of thighs brown with cream spots. 
Description of holotype. — Head broader than body, wider than 
long; head width 45.8% SVL; snout rounded in dorsal view, truncate 
in lateral profile; snout short, eye-nostral distance 71.2% eye length; 
canthus rostralis moderately sharp, straight; loreal region concave, 
sloping abruptly to lips; lips not flared; nostrils directed laterally, 
weakly protuberant; interorbital space weakly furrowed, edges of 
frontoparietals bearing low crests; upper eyelid broad, its width 
140.6% IOD; supratympanic fold obscure, covering upper edge of 
tympanum; tympanum prominent, higher than long, its length 37.9% 
eye length; tympanum separated from eye by distance equal to 
width of tympanum; skin of head tuberculate; eyelid bearing several 
large tubercles, one at posterolateral corner greatly elongated; tuber- 
cles along canthus forming canthal ridge; tongue about as long as 
wide, weakly notched posteriorly, posterior l A free; choanae mod- 
26 
OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
erate-sized, not concealed by palatal shelf of maxillae; prevomerine 
dentigerous processes broad, arched, posterior to choanae, separated 
medially by % choanal width, extending laterally to lateral edge of 
choanae, bearing a row of 10-12 teeth per process. 
Skin of dorsum tuberculate, tubercles heterogeneous, and bearing 
short paravertebral and dorsolateral ridges, most prominent of which 
are paravertebral scapular ridges connected by a transverse fold 
(Fig. 13); tubercles on flanks less heterogeneous than those of 
dorsum; skin of upper surfaces of limbs bearing heterogeneous 
tubercles; skin of venter smooth; no dorsolateral folds (other than 
short ridges); no trace of discoidal folds; posterior surfaces of thighs 
about vent areolate, a pair of prominent tubercles posterolateral to 
anus; forearm bearing a series of ulnar tubercles which are not larger 
than tubercles on upper surface of forearm; two palmar tubercles, 
outer bifid and larger than inner; no supernumerary palmar tuber- 
cles; subarticular tubercles round, low, flat, simple; fingers lacking 
lateral fringes; digits swollen apically, bearing ill-defined, narrow 
pads ( perhaps due to length of preservation ) ; heel bearing a single, 
non-elongate tubercle; outer edge of tarsus bearing a series of 
tubercles, inner edge of tarsus lacking tubercles or folds; two meta- 
tarsal tubercles, inner laterally compressed, four times as long as 
Fig. 14. — Eleutherodactylus cerastes (AMNH 88062). Photo by Charles 
W. Myers. 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 27 
wide, about three times size of low, round outer metatarsal tubercle; 
no supernumerary plantar tubercles; subarticular tubercles longer 
than wide, non-conical, simple, smaller than those of fingers; toes 
lacking lateral fringes and webbing; toes bearing dilated pads at 
tips, pads wider than long, much larger than those of fingers; hind 
limbs relatively long, heel of adpressed limb reaches to between 
eye and tip of snout; shank 60.0% SVL. 
Coloration in preservative. — The dorsum is brown and the ridges 
and tubercles dark brown to black; labial bars are present but 
canthal and supratympanic stripes are not distinct; the limbs are 
barred with dark brown, the dark bars are narrower than the brown 
interspaces. The groin, lower flanks, and anteroventral surfaces of 
the thighs are boldly marked with brown bars and spots on a pale 
cream background; the posterior surfaces of the thighs are dark 
brown with cream spots. The venter and undersides of the limbs are 
cream with moderate to intense brown motting and reticulation. 
The throat is brown punctated with cream. 
Variation. — Males lack vocal sacs and slits and tend to have 
larger ears than do females. Other than the variation in ear size, the 
proportions do not suggest sexual dimorphism. Males are smaller 
than females and within my samples the only mature females are 
46 and 55.S mm SVL. A female (USNM 195785, the holotype) is an 
immature female 46.7 mm SVL. The immature females (small eggs 
and weakly convoluted oviducts) range in size from 35-46.7 mm 
SVL. Females with straight oviducts are 32.7 mm SVL or smaller. 
The three adult or subadult Colombian specimens ( KU 144992- 
94) have weakly-flared lips and weakly-sloping snouts (in lateral 
profile). They also differ from the Ecuadorian specimens in having 
more rounded warts on the body. These differences might be simply 
geographic variation, but the less "sharp" warts may be due to the 
vicissitudes of preservation. A juvenile from Colombia also has 
more rounded warts ( Fig. 14 ) . 
Measurements of the holotype. — In mm, SVL 46.7. shank 28.0, 
head width 21.4, head length 16.3, eyelid width 4.8, IOD 3.4, tym- 
panum length 2.5, eye length 6.6, eye-nostril distance 4.7 
Etymology. — Greek, kerastes, horned: in reference to the eyelid 
tubercle. 
Distribution. — Known from low to moderate elevations (ca. 500- 
1580 m) along the Pacific versant of Colombia and Ecuador (Fig. 
10). 
Remarks. — E. cerastes appears closely allied to E. biporcatus, dis- 
tributed to the north, and to E. cornutus, found at comparable (or 
slightly higher) elevations on the Amazonian versant of the Ecua- 
dorian Andes. The three are readily distinguished, even as juveniles, 
in that E. biporcatus lacks elongate eyelid tubercles (present in 
cerastes and cornutus) and E. cornutus has a prominent inner tarsal 
28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
fold and lateral fringes on the toes (lacking in biporcatus and 
cerastes ) . 
Eleutherodactylus cornutus (Jimenez de la Espada) 
Limnophys comutus: Jimenez de la Espada, 1870:60 [Type-locality — Rio Suno, 
San Jose de Moti, Napo Prov., Ecuador]. 
Limnophys napaeus: Jimenez de la Espada, 1870:60 [Type-locality — Rio Suno, 
San Jose de Moti, Napo Prov., Ecuador]. New synonymy. 
Strabomantis cornutus: Jimenez de la Espada, 1872:85. 
Hylodes cornutus: Boulenger, 1882:220. 
Ctenocranius cornutus: Melin, 1941:49. 
Lithodytes cornutus (part): Anderson, 1945:45. 
Eleutherodactylus cornutus cornutus (part): Rivero, 1961:55. 
Eleutherodactylus cornutus: Gorham, 1966:66. 
Diagnosis. — A large Eleutherodactylus (immature 9 54.0 mm SVL) with 
a broad head (HW/SVL = 48.8-56.4%); snout truncate in lateral profile; 
frontoparietals bearing prominent lateral crests ending in elevated bosses; crests 
not extending onto nasals; tympanum prominent, 2/5 to % eye length; prevo- 
merine dentigerous processes arch-like, extending laterally to outer edge of 
choanae; male lacking vocal sac and slits; ulnar tubercles prominent; two palmar 
tubercles, outer largest, bifid; fingers bearing weak lateral fringes; tips of 
fingers bearing narrow discs; first finger longer than second; skin of dorsum 
coarsely tuberculate; H shaped series of folds in center of back; posterolateral 
corner of upper eyelid bearing one or two elongated tubercles; skin of venter 
smooth; tarsus bearing fold along inner margin; two metatarsal tubercles, inner 
larger, not compressed, outer small but easily detected; no supernumerary 
plantar tubercles; toes bearing lateral fringes without basal webbing; toes 
bearing dilated discs, discs larger than those of fingers; posterior surface of 
thighs black with cream flecks; venter brown with diffuse white blotches. 
Description. — (Also see Fig. 15). The only descriptions of this 
species are the brief descriptions of Jimenez de la Espada (1870). 
To prevent further confusion, such as that of Boulenger ( 1882) and 
Rivero (1961), the species is here redescribed on the basis of two 
young males (BMNH 1913.11.1.57, KU 143350), a young female 
(KU 123448), and 36 juveniles. Head as broad as or broader than 
body, wider than long; head width 48.8-56.4% (x = 52.2) SVL; snout 
Fig. 15. — Eleutherodactylus sulcatus (left, KU 143350) and E. cornutus 
(right, KU 123586). Photos by William E. Duellman. 
BROAD-HEADED ELEUTHERODACTYLIXE FROGS 29 
round in dorsal view, short and truncate in lateral profile; eye length 
greater than eye-nostril distance; canthus rostralis sharp, concave, 
sloping gradually to lips, lips weakly flared; nostrils protuberant, 
directed laterally; tip of snout not bearing ridge or proboscis, not 
extending much beyond lower jaw; interorbital region concave, 
edges of frontoparietals enlarged; upper eyelid tuberculate, tubercles 
near posterolateral corner of eyelid greatly elongated (Fig. 1); 
cranial crests ending in bosses posteromedial to eyes, least prominent 
anteriorly; tympanum prominent, slightly higher than long; tym- 
panic region not vertical, weakly oblique due to flaring of maxillary 
arch; tympanum length 39.5-54.3% eye length in specimens 20-27 mm 
SVL, 55.5, 60.3, 66.7% in specimens 35.5, 43.0, and 54.0 mm SVL; 
supratympanic fold tuberculate; edge of lower jaw bearing a row of 
prominent tubercles; tongue small, round, weakly notched behind, 
posterior edge free; choanae small, teardrop-shaped, well within 
border of jaw when roof of mouth viewed from directly above, 
separated medially by distance equal to 6 times width of choana; 
prevomerine teeth in transverse series on prominent, triangular 
processes lying posterior to choanae; 8-9 teeth per process, processes 
in median contact; male lacking vocal sac and slits. 
Skin of dorsum bearing numerous tubercles and ridges in ap- 
parently constant though complex pattern; prominent interorbital 
tubercle; ciliary ridges lateral to cranial crests; dermal ridges extend 
posteriorly from terminis of cranial crest as scapular folds with a 
cross bar in scapular region; dorsolateral region bearing rows of 
tubercles; center of back from scapular "H" to vent bearing numer- 
ous short folds; limbs bearing oblique bars of tuberculated ridges 
( Fig. 16 ) ; ulnar region bearing series of prominent flap-like tuber- 
cles; heel tuberculate but no tubercle elongated; skin of throat, 
venter, underside of limbs smooth except just below vent where skin 
is areolate; flanks areolate; discoidal fold present, ending anterior to 
groin; shank 54.7-63.0% (x = 57.4) SVL. 
Fig. 16. — Color patterns of hind limbs of Eleuiherodactylus cornutus (left, 
KU 123448) and E. sulcatus (right, KU 123586). 
30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Two palmar tubercles, outer bilobed (fused median and outer); 
supernumerary palmar tubercles at base of digits, not extending 
onto digits; subarticular tubercles round, simple, weakly conical; 
fingers bearing thin lateral fringes; digits bearing pads with circum- 
ferential grooves, pads less than Vk times digit width below pad; 
first finger longer than second. 
Inner edge of tarsus bearing distinct fold % tarsus length; outer 
edge of tarsus bearing row of conical tubercles; inner metatarsal 
tubercle elongate, not compressed; outer metatarsal tubercle promi- 
nent, longer than wide, not elevated, % size of inner; plantar surface 
bearing a few minute supernumerary tubercles; subarticular tu- 
bercles like those of fingers but smaller; toes bearing lateral fringe 
and basal web, web not enclosing basal subarticular tubercle on any 
toe; toes bearing pads, pads 1% times width of digit below pad; outer 
edge of foot and fifth toe bearing tuberculate ridge; third toe longer 
than fifth toe. 
Coloration. — In preservative, brown with reddish-brown ridges 
and warts edged with black; limbs barred with narrow black bars 
about one-fourth width of brown interspaces; labial barring dark 
brown to black; posterior surface of thigh black with silvery or 
golden flecks (Fig. 16); venter heavily reticulated with brown on a 
white ground color; throat brown with fine cream mottling; under- 
sides of limbs brown with diffuse white blotching; flanks brown with 
diffuse white spots. 
In life, E. cornutus was described as follows (W. E. Duellman, 
field notes) : "Dorsum dark dull reddish brown; posterior surface of 
thigh black with bluish-white flecks. Venter dark brown and grayish- 
white. Throat reddish-brown. Feet orange tan. Iris dull bronze 
with black reticulation and a median horizontal brown streak" (20 
October 1971). "dorsum dull olive-brown with black and reddish- 
brown markings. Groin and ventral surfaces of legs black and white. 
Throat and belly brown and white. Posterior surface of thigh black 
with small white flecks. Iris tan with brown triangles. Tongue 
bright yellow." (3 August 1968). 
Distribution. — Known only from moderate elevations (500-2000 
m) along the eastern base of the Ecuadorian Andes (Fig. 12). 
Remarks. — Authors referring to E. cornutus fall into two cate- 
gories, those citing the name without specimens and those applying 
the name to other species. Dunn (1944) applied the name cornutus 
to what has become the holotype of Ambh/phrijnus ingeri. Andersson 
(1945), Cochran and Goin (1970), and Puvero (1961) applied the 
name cornutus to specimens of E. sulcatus. Boulenger ( 1882), Coch- 
ran and Goin (1970), and probably Peracca (1914), used the name 
for young Amblyphrynus ingeri. 
I regard Limnophys napaeus Jimenez de la Espada as a synonym 
of E. cornutus. Jimenez de la Espada ( 1870) distinguished cornutus 
and napaeus on the basis of tongue shape, the narrow separation of 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 31 
the prevomerine teeth, the longer eye tubercle of cornutus, and 
minor differences in coloration. Both were included in the genus 
Limnophys, which was characterized in part by the tuberculate eye- 
lid. Examination of the type of napaeus could resolve the question 
of its identity ( Jimenez de la Espada published several illustrations 
of cornutus in his 1875 work), but the types of both species are 
evidently lost (William E. Duellman was unable to find them when 
he studied Jimenez de la Espada's material in 1969). Jimenez de la 
Espada ( 1872 ) placed napaeus in the synonymy of biporcatus, and 
this action was followed by all subsequent authors. Geographically, 
it seems reasonable to argue that cornutus and napaeus are iden- 
tical, especially in view of Jimenez de la Espada's (1870) descrip- 
tion in which the skin of the dorsum was characterized as warty for 
the 50 mm SVL type of napaeus. Comparable sized biporcatus 
would be characterized as having smooth skin with scattered warts. 
Additionally, the arched prevomerine processes of napaeus agree 
with cornutus but not biporcatus (prevomerine processes triangular 
in outline). 
Eleutherodactylus maussi (Boettger) 
Hylodes maussi Boettger 1893:99 [Type-locality. — Puerto Cabello, Estado Cara- 
bolo, Venezuela]. 
Eleutherodactylus maussi: Gines 1959:112. 
Eleutherodactylus cornutus maussi: Rivero 1961:54. 
Diagnosis. — A large Eleutherodactylus (adult 9 9 56-70 mm SVL) with a 
broad head ( HW/SVL = 44.3-54.9f); snout strongly sloping in lateral profile; 
lips flared in adults (flaring increases with size); frontoparietals bearing lateral 
crests, crests ending in moderate boss; crests not extending onto nasals; tym- 
panum prominent, 60-87% eye length; prevomerine dentigerous processes arch- 
like, extending laterally to outer edge of choanae; males with vocal sac and 
slits; ulnar tubercles prominent; fingers bearing weak lateral fringes; fingers 
bearing narrow discs at tips, first finger longer than second; skin of dorsum 
tuberculate; eyelid tuberculate, no tubercle elongate; skin of venter coarsely 
areolate; tarsus bearing fold along inner margin; two metatarsal tubercles, inner 
larger than outer, not compressed, outer prominent; plantar surface bearing 
numerous supernumerary tubercles; toes bearing lateral fringes but no basal 
webbing; toes bearing comparatively broad discs; posterior surface of thighs 
brown with cream flecks; venter cream. 
Descriptions. — Boettger (1893) and Nieden (1923) provide brief 
descriptions; that of Rivero (1961) is more complete. 
Distribution. — Known only from the Coastal Range of Venezuela 
(Fig. 10). 
Remarks. — E. maussi is certainly not conspecific with E. cornu- 
tus, but its relationship to E. sulcatus is close. The structural differ- 
ences between maussi and sulcatus are more extensive than Rivero 
(1961) realized. Heatwole (1962) reported on several aspects of 
the biology of maussi; comparable information is not available for 
any of the other broad-headed eleutherodactylines. 
32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Eleutherodactylus necerus new species 
Holotype. — USNM 195798, an adult female collected at Mindo, Pichincha 
Prov., Ecuador. 
Paratype. — USNM 195799, a juvenile female collected at the Rio Lelia, a 
tributary of the Rio Toachi, Pichincha Prov., Ecuador. 
Referred specimens. — Two juveniles: USNM 195852, Rio Blanco, near 
mouth of Rio Yambi, Pichincha Prov., Ecuador, 700 m; USNM 195851, road 
from Pacto to Rio Guayllabamba, Pichincha, Ecuador. 
Diagnosis. — A large Eleutherodactylus (adult female 93.3 mm SVL); snout 
truncate in lateral profile; lips not flared; frontoparietals bearing low parasagittal 
crests not ending in bosses, not extending onto nasals; tympanum % to 2/5 eye 
length in females; prevomers with broad, arched processes (adult only), ex- 
tending laterally to middle of choana ( not triangular ) ; ulnar tubercles present, 
not enlarged; two palmar tubercles, outer largest, bifid; fingers lacking lateral 
fringes; digit tips bearing discs, narrow; thumb longer than second finger; skin 
tuberculate and ridged above in young and adult frogs; upper eyelid tubercular, 
no tubercle elongate; skin of venter smooth; tarsus lacking tubercles or folds; 
two metatarsal tubercles, inner laterally compressed, outer obsolete, no super- 
numerary plantar tubercles; toes bearing lateral fringes, no basal webbing; toes 
with discs (broader than long), toe pads larger than those of fingers; posterior 
surfaces of thighs black with large cream spots; venter cream without brown 
mottling. 
Description of Jiolotype. — Head as broad as body; broader than 
long; head width 46.2% SVL; snout rounded in dorsal view, truncate 
in lateral profile; snout short, eye-nostril distance 78.4% eye length; 
canthus rostralis obtuse, concave; loreal region concave, sloping 
gradually to flared lips; nostrils directed laterally, weakly protuber- 
ant; interorbital region furrowed, narrow; upper eyelid 146.3% IOD; 
supratympanic fold present, not prominent, obscuring upper edge 
of tympanum; tympanum higher than long, its length 38.8% that of 
eye; tympanum separated from eye by 1/2-2 times width of tym- 
panum; skin of head smooth with numerous large warts; warts 
numerous and prominent on eyelid, none elongated (Fig. 13); can- 
thai folds present; tongue as long as wide, weakly notched posteri- 
orly, posterior % free; choanae large, not concealed by palatal shelf 
of maxillae; prevomerine dentigerous processes present, arched, 
posteromedial to choanae; processes in median contact, extending 
laterally to middle of choana; 13-16 teeth per process in an oblique 
row. 
Skin of dorsum and limbs bearing numerous small tubercles; 
dorsum bearing numerous short ridges, none elongate; transverse 
fold in scapular region; flanks not greatly tuberculate; skin of con- 
cealed surfaces and venter smooth; no enlarged post-anal tubercles. 
Forearm not bearing distinct ulnar tubercles or folds; two palmar 
tubercles, outer largest, bifid; no supernumerary palmar tubercles; 
subarticular tubercles round, non-conical, (not flattened) simple; 
fingers lacking lateral fringes except for a faint fringe on inner edge 
of second finger; tips of fingers swollen, bearing narrow, obscure 
discs; first finger much longer than second (tip of adpressed second 
finger reaches base of pad of thumb ) . 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 33 
Heel and tarsus lacking tubercles or folds; two metatarsal tuber- 
cles, inner laterally compressed, 4 times as long as wide, outer 
obscure, 1/6 to 1/7 the size of inner metatarsal tubercle; no super- 
numerary plantar tubercles; subarticular tubercles longer than wide, 
non-conical, simple; toes bearing thin lateral fringes and brief basal 
webbing (webbing does not encompass the proximal base of the 
basal subarticular tubercle); toes bearing discs, pads wider than 
long, much larger than those of fingers. 
Coloration (in preservative). — Dorsum medium to dark brown 
with indefinite darker brown spots and blotches; limbs barred with 
dark brown, bars about equal (or slightly narrower) in width to 
interspaces; venter cream or white except for some dark brown spots 
at edge of lower jaw and at base of arm; lower flanks bearing heavy 
black reticulation continuing posteriorly onto anterior surfaces of 
thigh, shank, and tarsus; posterior surfaces of thigh black with large 
cream spots; lower edge of dorsal pigmentation on concealed thighs 
and shanks is marked by heavy black reticulation which does not 
grade into white venter. 
Variation. — The paratype is a juvenile female (30.2 mm SVL). 
It differs from the holotype in having non-flared lips, heterogeneous 
warts on the dorsum, and in having a series of small tubercles along 
the outer edge of the tarsus. 
Measurements in mm. — Data for the holotype are presented first 
followed by the corresponding data for the paratype. SVL 93.3 
(30.2), shank 52.S (17.2), head width 43.1 (15.0), head length 
(12.5), eyelid width 9.0 ( ), interorbital distance 6.2 ( ), 
tympanum length 4.5 (1.7), eye length 11.6 (4.6), eye-nostril dis- 
tance 9.1 (3.4). 
Etymology. — Greek, nekeros, hornless, in reference to the ab- 
sence of an elongated tubercle on the eyelid. 
Distribution. — Known only from moderate elevations on the 
Pacific versant of Ecuador (Fig. 12). 
Remarks. — E. necerus is most similar to E. bufoniformis but 
differs in not having toe web enclosing the basal subarticular toe 
tubercles and in having arched rather than triangular prevomerine 
dentigerous processes. The two species are dichopatrically dis- 
tributed in the Chocoan region. 
Eleutherodactylus sulcatus (Cope) 
Hylodes sulcatus Cope, 1874:126 [Type-locality — Nauta, Depto. Loreto, Peru]. 
Hylodes macwcephalus Peracca, 1904:29 [Type-locality — Valle Santiago, Mo- 
rono-Santiago Prow, Ecuador], [new synonymy]. 
Ctenocranius koki Melin, 1941:45, fig. 27. [Type-locality — Taracua, Rio Uau- 
pes, Estado Amazonas, Brasil.] [new synonymy]. 
Lithodytes comutus (part): Andersson, 1945:45. 
Eleutherodactylus macwcephalus: Peters, 1955:348. 
Eleutherodactylus comutus comutus (part): Rivero, 1961:55. 
34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
[Eleutherodactylus] koki: Myers, 1962:198. 
Eleutherodactylus sulcatus: Gorham, 1966:103. 
Diagnosis. — A large Eleutherodactylus (adult females 28.8-59.7 mm SVL) 
with broad heads ( HW/SVL = 45.6-53.3%); snout sloping in lateral profile; 
lips flared in adults (degree of flaring increases ontogenetically); frontoparietals 
bearing prominent lateral crests, crests ending in a boss, crests usually not 
extending onto nasals; tympanum )i eye length in males, Yz to % eye length in 
females; prevomers with broad, arched processes, extending laterally to lateral 
edges of choanae; males lacking vocal sac and slits; ulnar tubercles present, 
prominent; two palmar tubercles, outer largest, bifid; fingers with weak lateral 
fringes; digit tips of fingers lacking discs; thumb longer than second finger; 
skin tuberculate above in young and adult individuals; upper eyelid tuberculate, 
occasionally an elongate tubercle present; skin of venter coarsely areolate; tarsus 
bearing fold along inner edge; two metatarsal tubercles, inner not greatly com- 
pressed, outer smaller, prominent; plantar surfaces bearing numerous super- 
numerary tubercles; toes bearing lateral fringes; tips of toes bearing discs; pads 
wider than long; posterior surface of thighs black with cream spots (Fig. 16); 
venter cream with some brown suffusion on throat and chest. The throat of 
males is darker than that of females. 
Descriptions. — (Fig. 13); Andersson (1945), Cochran and Goin 
(1970), Cope (1874), Melin (1941), and Peracca (1904). Those by 
Melin and Peracca are descriptions of immature specimens. The 
small size of males is striking; males have most of the ventral surface 
washed with dull brown. Comparable sized (as well as adult) 
females are cream below. The males examined do not have vocal 
slits or a vocal sac. In life, E. sulcatus was described as "dull tan 
with brown markings dorsally; posterior surface of thigh black with 
pale yellow flecks; venter creamy gray; iris dull gray." (W. E. 
Duellman field notes). 
Distribution. — Upper Amazon basin ( 150-950 m ) in western 
Brasil, eastern Ecuador, and eastern Peru (Fig. 10). Although no 
specimens are known, E. sulcatus must also occur in southeastern 
Colombia. 
Remarks. — The type specimens of koki, marcocephalus, and sul- 
catus agree in color pattern, snout shape, proportions, and tubercu- 
lation of the body, limbs, and hands and feet. Perraca (1904) 
separated macrocephalus and sulcatus on the basis of the former 
lacking frontoparietal crests and the associated interorbital furrow 
and because he thought macrocephalus differed from sulcatus in 
having an oblique orientation of the tympanum instead of a vertical 
orientation. The lectotype (here designated) of macrocephalus (the 
larger of the two individuals included in MZS 2930) is 35 mm in 
snout-vent length and that of sulcatus 47 mm SVL. The holotype of 
sulcatus does not have a vertically oriented tympanum as reported 
by Peracca ( 1904 ) ; all moderate to large-sized individuals of sul- 
catus have flared maxillae and hence an oblique orientation of the 
tympanum. As Peracca reported, the holotype of macrocephalus 
does not have frontoparietal crests and thus is not different from 
most individuals of its size class included in sulcatus. The fronto- 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 35 
parietal crests enlarge with increase in body size and their develop- 
ment occurs at smaller SVL in females than in males. The holotype 
of sulcatus is a female with exceptionally well developed crests com- 
pared to larger individuals and appears to represent an abnormal 
individual. 
Melin (1941) did not attempt to distinguish koki from sulcatus 
and did not cite distinguishing features. The holotype of koki is a 
juvenile and does not differ from immature specimens of sulcatus. 
In the absence of distinguishing features ( other than those unique to 
frontoparietal crest hypertrophy in the holotype of sulcatus), Cteno- 
cranius koki Melin and Hylocles macrocephalus Peracca are placed 
in the synonymy of Hylocles sulcatus Cope. 
Rivera's ( 1961 ) Eleutheroclactylus cornutus cornutus is the same 
as E. sulcatus. I agree with Rivera that the Coastal Range popula- 
tion of the sulcatus group (E. maussi) is related to an Upper Amazon 
population (E. sulcatus) but refrain from asserting a subspecific 
relationship in the absence of any evidence of integradation. The 
features distinguishing the two populations are more numerous than 
those cited by Rivero. 
There are several literature records of E. cornutus from Andean 
Colombia (Boulenger, 1882; Cochran and Goin, 1970; Dunn, 1944; 
and Peracca, 1914). All are apparently based on Amblyphrynus 
ingeri. 
Relationships of the Broad-Headed Eleutherodactylines 
Based on prior experience (subjective systematics) E. biporcatus, 
E. bufoniformis, E. cerastes, E. cornutus, and E. necerus seem to 
form a closely-knit group of species, and E. maussi and E. sulcatus 
form a second group. Within the first group, E. bufoniformis and 
E. necerus combine as an allopatric species-pair, and £. biporcatus, 
E. cerastes, and E. cornutus constitute a second Artenkreis. A third 
group of species includes A. helonotus and A. ingeri but the closeness 
of their relationships is comparable to that between E. cornutus and 
E. sulcatus, i.e., different species-groups within a genus. 
The question of relationships within the complex of broad-headed 
eleutherodactylines is of less interest than that of the relationships 
of members of this complex to frogs having "normal-width" heads. 
The commonality of superficial character-states between E. bufo- 
niformis, E. biporcatus, and a number of species in the fttzingeri- 
longirostris group has long been recognized. Cochran and Goin 
(1970) included E. bufoniformis in their Group I (= fitzingeri 
group auctorum). E. anomalus, E. fleischmanni, and E. taunts are 
species of the E. fitzingeri group that closely resemble E. bufoni- 
formis but have narrow heads and little or no development of 
cranial crests. These species, in turn, grade into the "smooth- 
skinned" members of Group I (e.g., E. achatinus, E. conspicillatus, 
36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
E. fttzingeri, E. w-nigrum among others). The relationships of 
Amblyphrynus are problematic. It might be a remnant of a "proto- 
eleutherodactylus" stock that had not acquired digital discs or the 
absence of digital discs in Amblyphrynus might be secondary. Dis- 
covery that the terminal phalanges have short lateral projections 
does not resolve the question. 
In an attempt to test my subjective concept of relationships, I 
re-evaluated the data from the viewpoint of cladistics and con- 
structed a cladogram following the methods described by Camin 
and Sokal (1965). The cladistic methodology results in a more 
objective base from which taxonomic decisions may be made. 
The greatest measure of subjectivity in a cladistic analysis is the 
determination of direction of change among character-states. Direc- 
tion of change for the majority of traits listed below is based on two 
assumptions: (1) Primitive proto-Eleutherodactylus had a morphol- 
ogy not unlike that exhibited by the extant genera Hylactophryne 
and Ischnocnema. The only change necessary to convert these frogs 
into Eleuiherodactylus is the acquisition of digital discs. (2) Char- 
acter-states sporadic in occurrence among frogs are more likely to be 
derived than primitive (e.g., eyelid tubercles, tarsal fold, super- 
numerary plantar tubercles, flared lips, cranial crests, and broad 
heads ) . 
On the basis of both assumptions, characteristic 5 is miscoded 
below. My initial cladistic run was made with the trait coded in 
the reverse of that listed below. The result was a lack of compatibili- 
ties with other traits and a high number of extra evolutionary steps. 
I recoded the trait (as listed below) and, with the discovery of 
compatibilities and a reduction in the number of extra steps, treated 
the feature as miscoded. Characteristic 6 is miscoded on the basis 
of assumption 1 (all five species of Hylactophryne and Ischnocnema 
have supernumerary plantar tubercles); however, on the basis of 
assumption 2, it is correctly coded ( supernumerary plantar tubercles 
are sporatic in occurrence). Fewer evolutionary steps are required 
when characteristic 6 is coded as listed below than when coded in 
the reverse. 
The characteristics (and their coding) used in the analysis are 
as follows: 
1. Prevomerine processes; triangular processes (0) are con- 
considered primitive to arched processes ( 1 ) . 
2. Texture of skin on venter; smooth venters ( ) are considered 
primitive to coarsely areolate venters ( 1 ) . 
3. Eyelid tubercles; the presence of elongated tubercles on the 
eyelid ( 1) is coded as derived, their absence as primitive (0). 
4. Tarsal fold; the presence of an inner tarsal fold is coded as 
derived ( 1 ) . 
5. Digital discs; the presence of discs on all fingers and toes is 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 37 
coded as primitive (0), the absence of discs on the fingers 
as derived ( 1 ) , and the absence of discs on all digits as 
further derived ( 2 ) . 
6. Supernumerary plantar tubercles; the presence of such tu- 
bercles in adults is coded as derived ( 1 ) . 
7. Lateral fringes on the toes; the absence of such fringes is 
coded as primitive ( ) , their presence as derived ( 1 ) , and 
their presence with basal (or more) webbing of the toes as 
further derived (2). 
8. Flared lips; flared lips are coded as derived (1). 
9. Snout shape; truncate snouts are coded as primitive (0), 
sloping snouts as derived ( 1 ) . 
10. Cranial crests; the presence of crests on the frontoparietals 
is coded as derived ( 1 ) . 
11. Head width; narrow heads (HW < 45% SVL) are coded as 
primitive (0), broad heads (HW > 45% SVL) as derived 
(1). 
12. Snout length; eye-nostril distance < eye length is coded as 
primitive (0), eye-nostril > eye length (long snout) is 
coded as derived ( 1 ) . 
The twelve characteristics and their 26 character-states were 
tabulated for each of twelve species of eleutherodactylines ( three of 
which are "narrow headed" taxa). The species are as follows: 
1. Amblyphrynus ingeri, 2. A. helonotus, 3. Eleiitherodactylus sul- 
catus, 4. E. maussi, 5. E. cornutus, 6. E. cerastes, 7. E. 1)iporcatus, 
8. E. bufoniformis, 9. E. necerus, 10. E. anomolus, 11. E. fitzingeri, 
and 12. E. conspicillatiis. A compatibility matrix (Table 1) was ob- 
Table 1. — Compatibility matrix ( 12 characteristics and 12 eleutherodac- 
tylixe frog species ) ; the patterns ( columns ) are the numbers of extra 
steps from each of 12 pattern tables. 
Patterns 
Compati- Extra 
Characteristics I II HI IV V VI VII VIII IX X XI XII bilities steps 
i. ~x I I I 2 I I I I o o o~~ 3 9 
2. 1X1111011000 4 7 
3. 11X110100000 6 5 
4. 111X21011000 4 8 
5. 2022X101100 5 9 
6. 00001X000000 10 1 
7. 111111X112 2 2 14 
8. 10 0X0000 10 1 
9. 00001000X000 10 1 
38 
OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
tained from the 12 pattern tables ( one for each characteristic ) and 
a cladogram of 22 steps was constructed by the monothetic method 
(Camin and Sokal, 1965) Were all characteristics parsimonious 
with one another, a cladogram of 14 steps would result. No trait was 
weighted more than another except by virtue of selection (Inger, 
1958). 
The resultant cladogram (Fig. 17) may be altered by adopting 
the equally parsimonious subcladograms (A' and B'). Choice of 
the alternative subcladograms will reflect characteristic bias on the 
part of the investigator. My biases are indicated by my choice of 
subcladogram A over A' and B over B'. A further bias is revealed 
with the assignment of taxonomic status to various clusters of 
OTUs. I have grouped OTUs 1 and 2 as the genus Ambhjphnjnus 
(primary trait — absence of digital discs) and assigned OTUs 3-12 to 
Eleutherodactijlus. The species of Eleutherodactyhis are further 
arranged into three clusters: OTUs 3 and 4 as the E. sulcatus group, 
OTUs 5-9 as the E. biporcatus group, and OTUs 10-12 as part of the 
E. fitzingeri group. Many more taxa belong to the E. fitzingeri 
group (E. longirostris group aucctorum) than the three species listed 
above but I know of no other nominate taxa (other than those as- 
signed to synonymies herein) of the E. biporcatus, E. sulcatus 
groups or of the genus Amblyphrynus. 
@<Q)® 
Fig. 17. — Cladogram depicting relationships among the hroad-headed eleu- 
therodactylines and three narrow-headed Eleutherodactyhis. Subcladograms 
A' and B' may be substituted at nodes A and B, respectively, resulting in 
equally parsimonious cladograms. Character-state shifts are indicated by hori- 
zontal bars and are numbered (see list of characteristics). 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 39 
Discussion 
The presence of simple digits (no pads or discs) in Amblyphry- 
nus led me to believe that these frogs were more primitive than 
EleutJwrodactylus (discs uniform). It is difficult to distinguish be- 
tween primitively simple digits and secondarily simple digits on the 
basis of digits alone. By not weighing traits and by simultaneously 
considering several traits I hoped that Camin and Sokal's technique 
would allow resolution of this dilemma. When I first tabulated the 
character-tables and compatibility matrix, I coded characteristic 5 
in reverse to that given above (first coding: no discs — 0, discs on 
toes only — 1, discs on all digits — 2). The resulting compatibility 
matrix with the lack of compatibilities and the high number of extra 
evolutionary steps suggested miscoding of this trait. In the clado- 
gram constructed from this initial analysis, the species of Ambly- 
phrynus remained derived taxa rather than primitive taxa in spite 
of an apparent miscoding of trait 5. I am now convinced that the 
simple digits of Amblyphrynus are secondarily simple as opposed to 
the primitively simple digits in many telmatobiine leptodactylids. 
I interpret the secondarily simple digits as an adaptation to terrestrial 
existence. For the most part, the species of the E. fitzingeri group 
are found on low vegetation in the herbaceous layer of the forests 
( within 1-2 m of the ground ) . The broad-headed taxa are found on 
the forest floor ( in litter by day; actively hopping about by night ) . 
The broad-headed ceratophryine leptodactylids (especially Cer- 
atophrys) are cannibalistic, and the width of their gape is probably 
of importance in capturing and eating other frogs (Cochran, 1961; 
Noble, 1931). Many other frogs, having normal head widths are 
frequently cannibalistic (e.g., Rana catesbeiana) but frog-eating 
habits are most often attributed to those frogs having large, broad- 
heads (Pyxicephalus adspersa of Africa is well-known in this re- 
gard). I examined the stomach contents of a few examples of 
broad-headed eleutherodactylines but found no frog remains. The 
frogs examined (and stomach contents) included one E. cerastes 
(large millipeds, spiders), one E. necerus (small to large beetles, 
large spider), and three E. sulcatus (small to large beetles). The 
only notable point concerning the stomach contents is the relatively 
large size of the prey (relative to the size of the frogs). If we 
accept Hutchinson's ( 1959 ) argument of comparing trophic features 
(gape here), greater head width seems to be a means of affecting 
niche separation without becoming a much larger frog. However, 
quantitative data on prey size and the frequency of frog-eating by 
most anuran species in nature remains to be investigated. 
The broad-headed eleutherodactylines are distributed largely 
allopatrically to the ceratophryine leptodactylids. Three species of 
Ceratophrys occur in northwestern South America. Ceratophrys 
stohmanni occurs in arid to semi-arid habitats in northwestern Peru 
40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
and southwestern Ecuador (Peters, 1967), just south of the distribu- 
tion areas of the Chocoan elements of the broad-headed eleuthero- 
dactylines (E. biporcatus, E. bufoniformis, E. cerastes, and E. ne- 
cerus). Ceratophrys cornuta is widely distributed in the Amazon 
drainage, ranging westward to the base of the Andes in at least 
Ecuador, where it occurs syntopically with E. sulcatus. Its distribu- 
tional area extends much farther to the east than does that of E. 
sulcatus, and it presently appears that E. sulcatus is much more 
abundant in Ecuador than is C. cornuta, which is best described as 
rare in Ecuador. To the south and east, C. cornuta is evidently more 
abundant, as judged from the length of series collected in those area 
contrasted to the widely disperse ( spatial and temporal ) collections 
of the species in Ecuador. Ceratophrys calcarata occurs in the Santa 
Marta region of Colombia and has also been reported from the 
upper Orinoco (Rivero, 1961) in Venezuela. Both of these areas are 
allopatric to the distribution areas of the several broad-headed eleu- 
therodactylines. 
Eleutherodactylus maussi of the Coastal Range of Venezuela is 
widely separated geographically from the remaining species of 
broad-headed eleutherodactylines. Eleutherodactylus cornutus and 
E. sulcatus are parapatrically distributed in eastern Ecuador (cor- 
nutus at higher elevations) and E. sulcatus ranges south into central 
Peru and east to western Brasil. Amblyphrynus ingeri occurs at 
elevations of 2000 m in Andean Colombia but is not sympatric with 
any other broad-headed eleutherodactyline. The Pacific versant 
and lowlands of Colombia and northern Ecuador contain the richest 
fauna of broad-headed taxa. Data are rather sketchy at present but 
E. cerastes appears to occur at slightly higher elevations along the 
versant than do the other four species in the region, but co-occurs 
with E. biporcatus in Colombia and E. necerus and A. helonotus in 
Ecuador. Two species (E. biporcatus and E, bufoniformis) are more 
clearly low elevation species than are the three intermediate eleva- 
tion taxa (A. helonotus, E. cerastes, E. necerus), but the several spe- 
cies may prove to be sympatric when more collecting has been done 
on the western Andean slopes. Although they may prove to be more 
generally sympatric, they are probably ecologically segregated in 
that E. bufoniformis, E. necerus, and A. helonotus have lateral 
fringes on the toes and are likely to occur in greater proximity to 
streams than E. biporcatus and E. cerastes (species lacking toe 
fringes ) . The two species of Eleutherodactylus with toe fringes are 
dichopatrically distributed (E. bufoniformis in Panama and north- 
ern Chocoan Colombia; E. necerus in northern Chocoan Ecuador) 
as are the two species lacking toe fringes (E. biporcatus ranging 
from Honduras to northern Chocoan Colombia; E. cerastes from 
central Chocoan Colombia to Ecuador). The allopatric distributions 
of these species pairs is suggestive of recent separation, ecologic 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 41 
complimentarity, and may reflect the Pleistocene contractions of 
forested habitats into forest islands ( Haffer, 1969 ) , now represented 
as "wet" islands in the lowland, humid forests. 
Several authors have commented on the generic status of the 
broad-headed eleutherodactylines (e.g. Myers, 1962; Rivero, 1961). 
The nominate genera having as type-species one of the members of 
this complex include, in addition to AmbJijphnjnus, Ctenocranius 
Melin (type-species by subsequent designation, Myers, 1962, Lim- 
nophys cornutus Jimenez de la Espada), Limnophys Jimenez de la 
Espada (type-species by subsequent designation, Myers, 1962, L. 
cornutus Jimenez de la Espada), and Strabomantis Peters (type- 
species by monotypy, S. biporcatus Peters). All of the generic names 
excepting Amblyphrynus are based on taxa in the biporcatus group. 
No generic name is available for the sulcatus group although Melin 
may have intended Ctenocranius for a sulcatus-type frog rather 
than the biporcatus-hog designated the tvpe-species by Myers 
(1962). 
The broad-headed eleutherodactylines with cranial crests have 
constituted a putative species-group or genus since 1864. The con- 
dition of head width is diagnostic but the presence of cranial crests, 
while characteristic, is not. These frogs tend to have narrow digit 
tips (i.e., lack pads), especially on the hands, have prominent tym- 
pana, and reach comparatively large size. Except for head width, 
these traits characterize but do not diagnose the putative group. To 
recognize a single and separate genus for the broad-headed eleu- 
therodactylines requires that the genus be founded on the basis of a 
single trait — head width — and ignores the variation in characteristics 
within the group. Furthermore, the broad-headed condition is 
approached by two species of "normal" head width Eleutherodac- 
tylus — E. anomalus and E. laticeps. Cladisitically (Fig. 17), the 
E. sulcatus group and Amblyphrynus could be united as a generic 
group contrasting with the more heterogeneous E. biporcatus group, 
but the distinctions between the two units (OTUs 1-4 and OTUs 
5-9) are few. In keeping with my generally conservative approach 
to genera (Lynch, 1971) and the use of derived character-states 
(loss of digital discs in Amblyphrynus), I continue to include the 
biporcatus and sulcatus groups in Eleutlierodactylus. 
Specimens Examined (From South America) 
Amblyphrynus hclonotus (2). Ecuador, Prov. Pichincha: Mindo, USNM 
195784; Rio Pitzara, BMNH 1970.178. 
Amblyphrynus ingeri (10). Colombia, Depto. Antioquia: El Clara Creek, 
AMNH 39978-80; Medellin, AMNH 38639, 38649. Depto. Cundinamarca: 
Bogota, BMNH 69.7.25.11; 8 km S Gachala, San Lsidro, FMNH 81915. Depto. 
Magdalena: west side of Sierra Nevada, Santa Marta, MCZ 17577. Depto. 
Tolima: Guindio mountains, MCZ 8237-38. 
Eleuthewdactyhis biporcatus (21): Colombia. Depto. Caldas: Pueblo 
42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
Rico, Santa Cecilia, 800 m, FMNH 54591. Depto. Choco: Alto del Buey, north 
slope, 300-1070 m, LACM 73154-57, 73177-80; Camino de Yupe, 350-625 m, 
LACM 73153, 73167-76. Depto. Valle: Raposa River Station, Rio Raposa, 
USNM 151447, UMMZ 132814. 
Eleutherodactylus bufoniformis (6). Colombia, Depto. Choco: Camino 
de Yupe, 350-500 m, LACM 73245-46; upper Rio del Valle, 50 m, LACM 
73247; Alto del Buey, north slope, 300-420 m, LACM 73248-49; Depto. Valle: 
Buenaventura, BMNH 95.11.16.54/1947.2.15.68 (holotype). 
Eleutherodactylus cerastes (20). Colombia, Depto. Cauca: Pueblo Rico, 
1580 m, BM 1910.7.11.61-62; El Tambo, La Costa, 1000 m, KU 144992-93; El 
Tambo, Munchique, 1000 m, KU 144994. Depto. Valle: 13 km W Dagua, Rio 
Anchicaya drainage, 850-1200 m, AMNH 88062. 
Ecuador. Imbabura Prov.: Lita, Rio Mira USNM 195786-88, 195795. 
Pichincha Prov.: Pachijal, USNM 195789-93; Palma Real, 195794; Rio Pitzara, 
USNM 195796(3); road from Pacto to Rio Guayllabamba, USNM 195797. 
Eleutherodactylus cornutus (43). Ecuador. USNM 167431. Prov. Morona- 
Santiago: Cutucu mountains, east of Macas, 1800-2000 m, AMNH 39499; 
Macuma, USNM 167401-08; Miazal, 167409-27. Prov. Napo: S slope Cordillera 
del Due above Rio Coca, 1150 m, KU 123447-48; Loreto, USNM 167428; Rio 
Azuela, 1740 m, KU 143350; Cerro Sumaco, AMNH 22265-70; S slope Cerro 
Sumaco, USNM 167429. Prov, Pastaza: Canelos, USNM 167430. Prov. Tun- 
gurahua: El Topo, 1270 m, BM 1912.11.1.57; Mirador, Banos, 1500 m, BM 
94.10.30.44. 
Eleutherodactylus maussi (46). Venezuela, Estado Aragua: Rancho 
Grande, AMNH 70533-52, 70554-58, MCZ 50701-02, UMMZ 113932(5), 
113933-34, 113935(2), 113936-41, 122376-77. Estado Carabolo: Puerto Ca- 
bello, MCZ 10178 (cotype); San Esteban, UMMZ 56474. 
Eleutherodactylus nccerus (4). Ecuador, Prov. Pichincha: Mindo, USNM 
195798; road from Pacto to Rio Guayllabamba, USNM 195851; Rio Blanco, near 
mouth Rio Yambi, USNM 195852; Rio Lelia, tributary of Rio Toachi, USNM 
195799. 
Eleutherodactylus sulcatus (93). Brasil, Estado Amazonas: Taracua 
GNM 494 (holotype Ctenocranius koki). Ecuador, Prov. Morona-Santiago: 
Miazal, USNM 167452-60; Valle Santiago, MZS 2930(2) (cotypes, larger here 
designated lectotype, Hylodes macrocephalus) . Prov. Napo: Limon Cocha, 
300 m, LACM 72161-62,UIMNH 55796; Loreto, USNM 167433-44; Payamino, 
USNM 167451; Puerto Libre, 570 m, KU 123585-87; Rio Cotopino, USNM 
167432; upper Rio Napo, 500 m, UMMZ 92111(2), 92114; Rio Suno, UMMZ 
92112; San Jose, Abayo Oriente, AMNH 22176, 22181, 22184; Santa Cecilia, 
340 m, AUM 7850-54, KU 108989, 123574-84; S slope Cerro Sumaco, USNM 
167445-50. Prov. Pastaza: Canelos to Marafion, Rio Bobonaza, MCZ 19640-41, 
USNM 139897; Don Tomas, 5 km S Montalvo, USNM 167462; Pacayacu, 
USNM 167466-68; region of Puyo, USNM 167469-70; 9 km SE Puyo, UMMZ 
126820; Rio Arajuno headwaters, USNM 167464; Rio Capahuari headwaters, 
USNM 167463; Rio Pindo, near town of Rio Tigre, USNM 167461; mouth of 
Rio Shiona at Rio Conambo, USNM 167465. Peru, Depto. Amazonas: Rio 
Cenipa, Ayendama, AMNH 42272; mouth of Rio Santiago, AMNH 42304. 
Depto. Junin: Chanchamayo, AMNH 42894. Depto. Loreto: Centre Union, 
TCWC 41699, 41738; Iquitos, AMNH 42293; Moropon, TCWC 39215-16; 
Manseriche range, head of Rio Caterpisa, 1500 ft., AMNH 42046; Nauta, ANSP 
11385 (holotype, Hylodes sulcatus); upper Rio Cushabatay, AMNH 42922; 
Rio Sapo, Rio Portongo, AMNH 43391; Rio Tampo (Tambo), AMNH 42091- 
92; upper Rio Utoquinea, AMNH 43136, 43377. Depto. San Martin: Rio 
Huallaga, Achinamisa, AMNH 42503. 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 43 
Summary 
The broad-headed eleutherodactyline frogs (head width 45-65% 
snout- vent length) occur in forested habitats from Honduras to 
Venezuela and Peru. Nine species in two genera are recognized. 
Seven species are placed in two species groups of Eleutherodactylus. 
The biporcatus group includes five species: biporcatus, bufoni- 
formis, cerastes sp. nov., cornutus, and necerus sp. nov. The sulcatus 
group includes two species: maussi and sulcatus. Two species of 
Ambtyphrynus are recognized: helonotus sp. nov. and ingeri. 
Based on an analysis of the characteristics evident among the 
nine species, the relationships of the taxa are examined cladistically. 
The cladistic analysis suggests that this complex of frogs is derived 
from frogs of the fitzingeri group. The discless frogs of the genus 
Ambtyphrynus are advanced representatives of the broad-headed 
group and differ from Eleutherodactylus in having lost digital discs. 
If the broad-headed frogs with digital discs are accorded generic 
recognition, the generic name would be Strabonwntis. 
Most published records of E. cornutus apply to Ambtyphrynus 
ingeri (Colombian records) or E. sulcatus (Ecuador, Peru). Limno- 
phys napaeus Jimenez de la Espada is removed from the synonymy 
of E. biporcatus and placed in the synonymy of E. cornutus. Hy- 
lodes macrocephalus Peracca and Ctenocranius koki Melin are as- 
signed to the synonymy of E. sulcatus. 
Three new species are named. Ambtyphrynus helonotus from the 
Pacific Andean versant of Ecuador differs from A. ingeri in lacking 
osteoderms, lacking a squamosal-maxillary connection, having arch- 
like prevomerine odontophores instead of triangular processes, lower 
cranial crests, no tarsal fold, and in coloration and proportions. 
Eleutherodactylus cerastes from the Pacific Andean versant of Co- 
lombia and Ecuador differs from E. biporcatus in having an elongate 
eyelid tubercle, arch-like prevomerine processes, and smaller tym- 
pana; cerastes differs from cornutus in lacking tarsal folds and lateral 
fringes on the toes and in having pale spots of the concealed thigh 
surfaces. Eleutherodactylus necerus from the Pacific Andean versant 
of Ecuador differs from E. bufoniformis in lacking toe webbing, 
having arch-like prevomerine processes, and in lacking brown 
mottling on the venter. 
Broad-headed eleutherodactyline species are primarily para- 
patrically distributed among themselves as well as relative to other 
broad-headed leptodactylids, especially Ceratophrys. Eleutherodac- 
tylus biporcatus and E. bufoniformis are sympatric over much of 
Chocoan Colombia and forested Panama. Ambtyphrynus helonotus, 
Eleutherodactylus cerastes, and E. necerus may prove to be sym- 
patric on the Andean slopes of western Ecuador. Between sub- 
families ( Ceratophryinae and Telmatobiinae), sympatric occurrence 
of Ceratophrys cornuta and Eleutherodactylus sulcatus constitutes 
44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 
the only contradiction of this generalization. The largely parapatric 
pattern of distribution is suggestive of potential competition among 
broad-headed leptodactylids. 
Resumen 
Las ranas eleutherodactylinas con cabecas ancha (ancha de la 
cabeza equal 45-65 porcent hocico-ano longitud) viven en bosques 
entra Honduras hasta Venezuela y el Peru. Nueve especies de dos 
generos son reconocadas. Siete especies son colocadas en dos espe- 
cies-grupos del genero Eleutlierodactylus. El grupo biporcatus 
contiene cinco especies: biporcatus, bufoniformis, cerastes esp. nov., 
cornutus, y necerus esp. nov. El grupo sulcatus contiene dos espe- 
cies: maussi y sulcatus. Dos especies del genero Amblyphrynus 
son reconocadas — helonotus esp. nov. y ingeri. 
Las afinidades de los nueve especies son examinadas cladistica- 
mente. Es analisis se indica que esta collection de ranas se derivadas 
de ranas del grupo ftzingeri (genero Eleutherodactijlus) . Las ranas 
que faltan discos (genero Amblyphrynus) son avanzado especies de 
los grupos con ancha cabecas y son distinto de Eleutherodactijlus 
porque ellos faltan todos discos de los dedos. Si las ranas con 
cabecas ancha y discos de los dedos se acomodado reconocimiento 
generico, el nombre generico estara Strabomantis (en lugar de dos 
especies-grupos de Eleutherodacti/lus) . 
La parte mayor de las anotaciones publicadas de E. cornutus son 
actualamente Amply phrymis ingeri (anotaciones colombianos) o E. 
sulcatus (anotaciones ecuatorianos y peruanos). Limnophys na- 
paeus no es un sinonimo de E. biporcatus pero es un sinonimo de 
E. cornutus. Hylodes macrocephalus y Ctenocranius koki son colo- 
cadas en la sinonimia de E. sulcatus. 
Tres especies nuevas se llaman. Amblyphrynus helonotus de las 
faldas andinas pacifico del Ecuador se distinguido de A. ingeri en 
faltando osteodermos, faltando un articulation escamoso-maxilar, en 
teniendo odontoforos prevomerinos arco-semejante en lugar de pro- 
tuberancias triangular, crestas del craneo mas bajo, no plieque tarsal, 
y en coloration y proporciones. Eleutherodactijlus cerastes de las 
faldas andinas pacifico de Colombia y Ecuador se distinguido de 
E. biporcatus en teniendo un tuberculo parpado bien alargado, 
protuberancias prevomerinos arco-semejante, y un timpano menor; 
cerastes se distinguido de cornutus en faltando plieques tarsal y 
orillas laterales en los dedos (pes), y en teniendo puntos palidos en 
los superficies del muslo callado. Eleutherodactijlus necerus de las 
faldas andinas pacifico de Ecuador se distinguido de E. bufoniformis 
en faltando membranas interdigital de pes, en teniendo odontoforos 
prevomerinos arco-semejante en lugar de protuberancias triangular, 
y en teniendo un abdomen bianco (en lugar de moteado con 
moreno ) . 
BROAD-HEADED ELEUTHERODACTYLINE FROGS 45 
En general, los especies eleutherodactylinos con cabecas ancha 
tienen distributions parapatricos entre ellos mismos y tambien 
relative) a otra leptodactylidos con cabecas ancha, especialmente 
Ceratophrys. Eleutherodactylus biporcatus y E. bufoniformis son 
macrosimpatrico en Colombia occidental y Panama. Amblyphrynus 
helonotus, Eleutherodactylus cerastes, y E. necerus son encontrado 
tal vez simpatricamente en las faldas andinas pacifico en Ecuador. 
Entre los subfamilias ( Ceratophryinae y Telmatobiinae ) , suceso 
simpatrico de Ceratophrys cornuta y Eleutherodactylus sulcatus 
constitye el contradiction solo de este generalizacion. El modelo 
de distribution (principalmente parapatrico) es sugestivo de com- 
petition potential entre los leptodactylidos con cabecas anchas. 
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