Occasional papers of the Museum of Natural History, the University of Kansas

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

MCZ
LIBRARY

OCCASIONAL PAPERS ^ftf g \ 1989

of the

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 129, PAGES 1-18 APRIL 19,1989

A NEW GENUS AND SPECIES OF TOAD

(ANURA: BUFONIDAE) WITH AN

EXTRAORDINARY STREAM-ADAPTED TADPOLE

FROM NORTHERN HONDURAS

By

James R. McCranie1, Larry David Wilson2, and
Kenneth L. Williams3

In early June, 1980, at the end of the dry season, JRM and LDW travelled
to the Cordillera de Nombre de Dios south of the port of La Ceiba, Atlantida,
Honduras. Our objective was to sample the herpetofauna of the highest
elevations in the region of Cerro Biifalo. While working in cloud forest in
these steep mountains at 1760 m, we found a single apparently undescribed
female bufonid. On the return trip, we collected a second specimen, a male,
in highland rainforest at 940 m and a series of peculiar tadpoles which we
suspected belonged to the same taxon.

In August of 1982 we returned to this area in the rainy season to try to link
the strange tadpole and the undescribed toad. An additional adult male and
numerous juveniles and tadpoles were collected. One of the tadpoles was in
the process of metamorphosis and allowed its association with adults.

A third trip to the same area in August, 1984, netted one adult male and
more juveniles and tadpoles, including a number of metamorphosing indi-
viduals.

'10770 S.W. 164th Street, Miami, Florida 33157.

department of Biology, Miami -Dade Community College, South Campus, Miami, Florida
33176.

'Department of Biological Sciences, Northwestern State University of Louisiana, Natchi-
toches, Louisiana 71457.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

The larva of this new toad is a mountain stream inhabitant and appears to
be the fourth example of a gastromyzophorous tadpole (Inger, 1966). This
terminology describes the condition of a well-developed, extensive ventral
suctorial disc anastomosed to the larval mouthparts, an adaptation for
clinging to smooth rock surfaces in swift mountain streams. The condition has
been reported in three genera of anurans belonging to two families and
occurring in both hemispheres. The bufonid genus Atelopus occurs in humid
forests in areas of some relief from Costa Rica southward to northern Bolivia
(Frost, 1985). The ranid genus Amolops occurs in mountain streams from
"Nepal, northeastern India, western and southern China to the Greater Sunda
Is." (Frost, 1985: 452). The third is the ranid Rana sauteri from Taiwan
(Kuramoto et al., 1984). However, the tadpole of/?, sauteri differs noticeably
from the other gastromyzophorous tadpoles in having a less well-developed
abdominal disc.

The tadpole of the new toad bears a striking resemblance to those known
for the species of Atelopus. The resemblance is so close that were it not for the
known association of the tadpole and adult toad, it would be tempting to
regard the tadpole as the larva of an undescribed species of Atelopus. In light
of the distinctiveness of the Cerro Bufalo toads (see diagnosis and description
below), we refer them to a new bufonid genus.

MATERIALS AND METHODS

Data on myology and osteology of the new genus were taken from a single
male (KU 206733) that was first dissected and then alizarin-cleared and
stained. Thigh musculature nomenclature and features used herein were
described in detail by Grandison (1978) and jaw musculature by Starrett
(1968). Bone terminology follows Trueb (1973) except for the shape of the
terminal phalanges (after Lynch, 1971) and the nature of the occipital groove
and orbitosphenoid, which follow McDiarmid (1971) and Cannatella (1986),
respectively. Measurements taken were defined by Duellman (1970) and
webbing formula by Myers and Duellman (1982). Tadpoles were staged
according to Gosner's (1960) system. Data on other bufonid genera were
taken from the literature with a few exceptions (see introduction to Compari-
sons section).

SYSTEMATICS

Atelophryniscus gen. n.

Type species. Atelophryniscus chrysophorus new species.

Diagnosis. A small bufonid distinguished from other genera in the
family by the following combination of characters: (1) habitus bufoniform;
(2) dorsal surface rugose, covered with scattered tubercles; (3) parotoid

A NEW GENUS AND SPECIES OF HONDURAN TOAD 3

glands distinct, subtriangular; (4) cranial crests low; (5) tympanic annulus
present, concealed by skin; (6) columella present; (7) ostia pharyngea present;
(8) single sinistral vocal slit present in adult males; (9)modal toe webbing
formula I 1-1 II 1-2 III 1-272 IV 3-1 V, subarticular tubercles small,
flattened; (10) outer metatarsal tubercle small, ovoid; (11) inner tarsal fold
present; (12) phalangeal formula of hands 2-2-3-3, feet 2-2-3-4-3; (13) adult
males 33.6-37.4 mm (n=2) snout-vent length, single female 36.2 mm; (14)
testes slender, cream colored, 76 snout- vent length; (15) adductor longus
muscle absent; (16) tensor fasciae latae short, arising from distal 73 of ilium,
inserting onto cruralis muscle at proximal 74 of thigh; (17) iliacus externus
muscle bulky, its origin at about mid-length of ilial shaft; (18) broad origin to
dorsal portion of semitendinosus; (19) adductor mandibulae externus super-
ficialis and posterior subexternus present, in S and E condition; (20) eight
presacral vertebrae; (21) neural arches imbricate; (22) cervical not fused to
second vertebra; (23) occipital condyles and cervical cotyles closely juxta-
posed; (24) no intervertebral fusion; (25) pectoral girdle fully arciferal; (26)
omosternum absent; (27) sacral-coccygeal articulation bicondylar; (28) coccyx
with a very small posteriorly-directed transverse process on left side only,
without expanded lateral edges; (29) terminal phalanges of hands expanded,
moderately T-shaped, those of feet knobbed; (30) two centralia in hand; (31)
frontoparietal bones not fused on midline; (32) frontoparietal fontanelle
obscured by medial articulation of frontoparietals; (33) occipital groove
open; (34) orbitosphenoid unossified; (35) otic ramus of squamosal extend-
ing over prootic; (36) palatine bone present, well-developed; (37) quadratoju-
gal in bony contact with maxilla; (38) larvae gastromyzophorous.

Etymology. The generic name is derived from the Greek prefix a-,
meaning "without," the Greek adjective telo, meaning "complete," the Greek
noun phryne, meaning "toad," and the Greek diminutive suffix -iscus. The
name is a reference to the resemblance of the tadpole to that of species of
Atelopus and the resemblance of the adult to species of Bufo, as well as to the
small size of the adults. The gender is masculine.

Atelophryniscus chrysophorus sp. n.
(Fig. 1)

Holotype. Adult male, University of Kansas Museum of Natural History
(KU) 206730, from Quebrada de Oro (15° 38'N, 86° 47'W), elevation 880 m,
tributary of Rio Viejo, south slope of Cerro Biifalo, Cordillera de Nombre de
Dios, Depto. Atlantida, Honduras, collected on 16 August 1982 by James R.
McCranie, Kenneth L. Williams, and Larry David Wilson.

Paratypes. KU 206731, adult female, from south slope of Cerro Biifalo,
elevation 1 760 m, above the Quebrada de Oro, Cordillera de Nombre de Dios,
Depto. Atlantida, Honduras, collected on 3 June 1980 by Gustavo A. Cruz

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

"

Fig. 1. Dorsal view of female paratype of Atelophryniscus chrysophorus (KU 206731).

Diaz, Mario R. Espinal, James R. McCranie, and Larry David Wilson; KU
206732, adult male, same data as holotype except elevation 785 m and
collected on 16 August 1984.

Description of holotype. Snout-vent length (SVL) 33.6 mm; tibia length
16.7 mm, 49.7 per cent of SVL; hand length 10.6 mm, 3 1.5 per cent of SVL;
footlength 15.3 mm,45.5percentofSVL; head length 10.6 mm, 3 1.5 per cent
of SVL; head width 11.3 mm, 33.6 per cent of SVL; snout in dorsal aspect
broadly acuminate, in profile bluntly rounded; snout length 2.5 mm, 23.6 per
cent of head length; distance from nostril to tip of snout 0.38 times less than
distance between nostril and eye; loreal region slightly concave; nostrils
protuberant, in line with anterior terminus of upper jaw, slightly below can thai
ridge; diameter of eye 4.6 mm, 43.4 per cent of head length; width of eyelid

A NEW GENUS AND SPECIES OF HONDURAN TOAD 5

3.6 mm, 31.9 per cent of head width; pupil horizontally elliptical; palpebral
membrane clear with a few silver flecks and a dusting of melanophores along
upper and lower edges; internarial distance 2.9 mm, 25.7 per cent of head
width; interorbital distance 4.1 mm, 36.3 per cent of head width; top of head
flat, slightly concave between nostrils, almost smooth; skin co-ossified with
skull; supraorbital crest low, pustulose, confluent with short parietal crest and
low postocular crest; supratympanic crest short, bulbous, in contact with
parotoid gland; upper surface of eyelids slightly pustulose; parotoid gland
subtriangular, slightly larger than upper eyelid, extending posteriorly to level
of axilla; tympanum concealed by skin; tongue lanceolate, barely free behind;
single sinistral vocal slit, at the posterolateral edge of tongue, extending back
to near angle of jaw; choanae subcylindrical, posterior to nostrils; prevomer
edentate; ostia pharyngea present, posteromedial to angle of jaw.

Arms slender, fingers long, slender; length of fingers from shortest to
longest 1-2-4-3; phalangeal formula of hand2-2-3-3; fingers freeof webbing;
subarticular tubercles on hands small, subcorneal; supernumerary tubercles
small, flattened; inner palmar tubercle round, slightly elevated; outer palmar
tubercle large, round, flat, about twice size of inner palmar tubercle; prepol-
lical protuberance present, with numerous tiny brown nuptial excrescences;
inner tarsal fold present; toes long, slender; webbing formula 1 1-1 II 1-2 III
1-272 IV 3-1 V; length of toes from shortest to longest 1-2-5-3-4; phalangeal
formula of foot 2-2-3-4-3; subarticular tubercles on toes small, flattened;
supernumerary tubercles on foot few, small; outer metatarsal tubercle small,
ovoid; inner metatarsal tubercle subtriangular, relatively small, about three
times the size of the outer metatarsal tubercle.

Cloacal opening directed posteriorly at upper level of thighs. Anal sheath
short. Skin of dorsum with moderate-sized scattered tubercles. Small scat-
tered tubercles present on upper surfaces of forelimbs and hindlimbs. Skin of
throat, chest, and belly pustulose.

Color (in life): dorsum pale lime green with olive brown blotching and
rusty-red-tipped pustules; pale orange pustules outlining lower edge of pale
lime green lateral stripe; lateral stripe bordered below by dark brown line
grading to olive brown ventrally; side of head pale yellow, mottled with dark
brown in front of and below eye; rusty-red dark brown-bordered bar extend-
ing from eye to posterior edge of lip; iris pale metallic green; limbs pale lime
green dorsally with olive brown blotches; toe tips and outer edge of web
orange; venter pale blue-green, motded with dark chocolate brown.

Color (in preservative): dorsum pale gray with grayish brown blotching;
pustules dull orange; venter cream, mottled with brown.

Variation in the paratypes. The male paratype (KU 206732) has a snout-
vent length of 37.4 mm, the female (KU 206731) 36.2 mm. Other measure-
ments and ratios are as follows (data for male paratype listed first): tibia length
18.2 mm, 17.9 mm, 48.7, 49.4 per centS VL; hand length 11. 5 mm, 12.2 mm,

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

30.7, 33.7 per cent S VL; foot length 17.2 mm, 17.0 mm, 46.0, 47.0 per cent
SVL; head length 12.3 mm, 11.6 mm, 32.9, 32.0 per cent SVL; head width
13.0 mm, 12.0 mm, 34.8, 33.1 per cent SVL; snout length 2.9 mm, 2.8 mm,
23.6, 24. 1 per cent head length; diameter of eye 4.6 mm, 4.6 mm, 37.4, 39.7
per cent head length; width of eyelid 3.8 mm, 3.6 mm, 29.2, 30.0 percent head
width; internarial distance 3.4 mm, 3.0 mm, 26.2, 25.0 per cent head width;
interorbital distance 4.5 mm, 4.5 mm, 34.6, 37.5 per cent head width.

The paratypes are similar morphologically to the holotype, except the
female lacks nuptial excrescences and a vocal slit.

Color (in life) of female (KU 206731): middorsum dark lime green,
flanked by pale lime green dorsolateral stripes; tubercles on dorsum brick red;
flanks dark lime green with red-tipped tubercles; chin mottled with brick red
and pale green; venter marbled with red-speckled gray and pale green; arms
and legs pale lime green with dark brownish green bands and brick-red-tipped
tubercles; iris lime green with some black reticulations.

Osteology. The premaxillae are separated medially, in contact anterome-
dially, slightly separated posteromedially. The pars palatina is edentate and
there is a well-developed pointed posteromedially-oriented palatine process
on each premaxilla. The quadratojugal is in bony contact with the maxilla.
The otic ramus of the squamosal extends over the prootic.

The cervical cotyles are closely juxtaposed (Type II of Lynch, 1971). The
neural arches are imbricate. There are eight presacral vertebrae. The widths
of the moderately long transverse processes from narrowest to widest are 2-
8-7-6-5-4-3. The transverse processes of the second presacral vertebra extend
laterally, those of 3, 4, and 5 extend slightly to moderately posterolateral^,
those of 6 and 7 extend laterally, and those of the eighth slightly anterolater-
ally. The sacral diapophyses are expanded. The sacral-coccygeal articulation
is bicondylar. The coccyx bears a small, posteriorly-directed transverse
process on the left side only, without expanded lateral edges. There is a well-
developed dorsal crest extending about three-fourths of the length of the
coccyx.

The pectoral girdle is fully arciferal and bears no omosternum. The
sternum is bony and bears no xiphisternum. The clavicle is bony.

The terminal phalanges of the hands are expanded and moderately T-
shaped, those of the feet are knobbed. There are two centralia in the hand.

Myology. The adductor longus muscle is absent in Atelophryniscus. The
tensor fasciae latae muscle is short, arising from the ventral surface of the
distal third of the ilium and inserting on the cruralis muscle at about the
proximal fourth of the thigh. The iliacus externus muscle is bulky, its origin
lies at about the midlength of the ilial shaft, its fibers arising mainly from the
dorsal and dorsomedial surface of the bone, with some additional fibers given
off from the lateroventral surface. There is a broad origin to the dorsal portion
of the semitendinosus. There is no accessory head to the adductor magnus

A NEW GENUS AND SPECIES OF HONDURAN TOAD 7

muscle. Both the adductor mandibulae extemus superficialis and posterior
subexternus are present and in the S and E condition.

Description of the tadpole (Figs. 2 and 3). A typical tadpole (in lot
number KU 206748) may be described as follows: developmental stage 37,
body length 9.6 mm, total length 26.4 mm; body depressed, flattened
ventrally, 1.5 times as wide as deep; snout broadly rounded in dorsal and
lateral profiles; eyes moderately large, separated by distance equal to about
three times diameter of eye, directed anterolaterally; nostrils small, consid-
erably closer to eye than tip of snout; spiracle sinistral, directed posterodor-
sally, located low on body, at about three-fourths of body length; anal tube
medial, long; caudal musculature deep at base, gradually tapering posteriorly
to about midpoint where tapering noticeably increases, extending nearly to tip
of rounded tail; dorsal fin slightly arched and deepest at about midpoint of tail;
ventral fin not noticeably arched; fins not extending onto body.

3 mm

Fig. 2. Lateral view of the tadpole of Atelophryniscus chrysophorus (KU 206748) in Gosner
stage 35.

Mouth ventral (Fig. 4), nearly as wide as body, followed by large suctorial
disc with raised rim; suctorial disc border complete laterally and posteriorly;
suctorial disc extending about three-fourths length of body, broadly rounded
posteriorly; median portion of upper and lower lip bare, rest bordered by one
row of small papillae, additional row present just inside outer row laterally on
upper labium; lateral folds well-developed; two upper and three lower rows
of denticles, all complete and about equal in length, extending to lateral edge

3 mm

Fig. 3. Dorsal view of the tadpole of Atelophryniscus chrysophorus (KU 206748) in Gosner
stage 35.

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 4. Mouthparts and ventral suctorial disc of the tadpole of Atelophryniscus chrysopho-
rus (KU 206748) in Gosner stage 35.

of mouth; beaks thin and without serrations, upper beak curved, slightly
longer than V-shaped lower beak.

Color in life: body black with gold markings; caudal musculature black
with pale yellow bands or spots; in preservative: body dark brown with tan
markings slightly evident; caudal musculature brown with pale tan bands or
spots; fins clear, lightly punctated with brown.

A tadpole in stage 42 (lot number KU 206748) has a body length of 10.8
mm and a total length of 25.4 mm. The beaks have disappeared and remnants
of the second upper denticle row are all that remain. The suctorial disc has
been partially resorbed and extends only slightly posterior to the axilla.
Minute red tubercles are present on the dorsum and forelimbs, red tubercles
on the hind limbs somewhat better developed. The webbing on the feet and
the banding of the hind limbs is similar to recently-metamorphosed individu-
als.

A juvenile (lot number KU 206748) with a tail stub is 9.2 mm in snout- vent
length; a juvenile (lot number KU 206748) with the tail completely resorbed

A NEW GENUS AND SPECIES OF HONDURAN TOAD 9

is 9.8 mm SVL. Color notes (in life) for these toadlets: olive green dorsum
with red-tipped pustules; pale yellowish green lateral stripes; dorsal spots red,
outlined with dark brown; bands on legs reddish brown, separated by pale
yellowish green; venter turquoise blue, mottled with chocolate brown; iris
metallic green.

Tadpoles in late stages of development, metamorphosing toadlets, and
juvenile toads were collected (KU 206734^9). Uniformity of toe webbing,
banding of the hind limbs, and red-tipped tubercles on the dorsum and hind
limbs of all these specimens and the adults unquestionably confirm the link
between the tadpoles and adults.

Etymology. The specific name chrysophorus is derived from the Greek
noun chrysus, meaning "gold," and the Greek suffix -phora, meaning
"bearing." The reference is to the gold markings on the dorsum of the
otherwise black tadpole, to the name of the stream at the type locality,
Quebrada de Oro (= "stream of gold"), and to the fact that the gold-spotted
tadpole occurred in that stream.

Natural history. The type locality, Quebrada de Oro, is a mountain stream
3-4 m wide. The stream has variously-sized boulders in the areas of rapids,
interspersed by broad, deep pools. The Quebrada de Oro flows into the Rio
Viejo, which joins the Rio Cangrejal, a major river of the mid-northern coast
of Honduras that flows through the port of La Ceiba on the Caribbean.

The vegetation of the area (at ca. 800 m) can be characterized as Subtropi-
cal Wet Forest (Holdridge, 1967) and less formally as highland rainforest.
Trees of the forest include sweetgum (Liquidamber), Cecropia, Ficus,
coralbean (Erythrina), oaks (Quercus), wild avocados (Persea), and Mi-
crotia. Some leguminous and rubiaceous trees and bactrid palms were also
present. The undergrowth consisted of dumbcane (Diefenbachia), various
small palms, Solatium sp., elephant ears, and other small aroids, Piper sp.,
Selaginella sp., and a few terrestrial bromeliads. Vines of the families
Solenaceae, Apocynaceae, Curcubitaceae, and Bignoniaceae were present, as
well as Passiflora, Mikania, Vitis, and Smilax. Vining aroids and ferns were
seen, as were epiphytes, including Spanish moss and other small bromeliads,
orchids, peperomias, ferns, aroids, mosses, and liverworts. No large, water-
filled epiphytic bromeliads or tree ferns were seen.

At higher elevations the highland rainforest along the Quebrada de Oro
gradually gives way to hardwood cloud forest (=Lower Montane Wet Forest
formation of Holdridge, 1967). The stream begins at about 1500 m; above this
point there is no contained water because of the steep slopes.

The adult female was found in a hole covered with vegetation at the base
of a low embankment in undisturbed hardwood cloud forest at 1760 m. All
other adults, juveniles, and tadpoles came from lower elevations (750-1090
m) in highland rainforest. The adult males came from within the forest along
the stream during the day or at dusk and the juveniles were found hopping on

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

the banks of the stream during the day or at night. No Atelophryniscus were
heard calling. The tadpoles usually inhabit the middle, fast-flowing portion
of the stream, in relatively deep pools, where they cling to the surface of
submerged boulders. When disturbed, they creep down the boulder, swim to
the bottom, or swim to another boulder. Tadpoles sometimes occur in
shallower areas where water moves rapidly over the rocks, or in side pools
with slower moving water. None were found in stagnant side pools. Subaerial
adhesion was occasionally observed in the Atelophryniscus tadpole, as has
been recorded for the gastromyzophorous larvae ofAmolops [Liu, 1950, (as
Staurois); Wassersug and Heyer, 1983].

COMPARISONS

Information in this section was taken largely from Cannatella (1986),
Grandison (1978, 1980a, 1981), Hoogmoed (1985), Inger (1966), Laurent
(1986), McDiarmid (1971), Pillai and Yazdani (1973), Pregill (1981), Ruiz-
C. and Hernandez-C. (1976), Savage and Kluge (1961), Tihen (1960), and
Trueb (1971), unless otherwise noted. Two colleagues (see Acknowledg-
ments) also provided unpublished information on various New World genera.

New World Genera. Currently there are twelve genera of bufonids
recognized in the New World (Frost, 1985; Hoogmoed, 1985; Cannatella,
1986; this paper). Cannatella (1986) divided ten of these genera into two
groups. The atelopodines Atelopus, Dendrophryniscus, Frostius, Melano-
phryniscus, Oreophrynella, and Osornophryne have partial or full fusion of
the epicoracoid cartilages, whereas Bufo, Crepidophryne, Peltophryne, and
Rhamphophryne have unfused epicoracoid cartilages (Cannatella, 1986).
The recently-described Andinophryne (Hoogmoed, 1985) exhibits the ate-
lopodine condition. Atelophryniscus displays the bufonine condition.

Atelophryniscus can be further distinguished from the "atelopodines" by
the following combination of characters: tensor fasciae latae short (elongate
in Atelopus, Frostius, and Osornophryne); adductor mandibulae in S+E
condition (S in Andinophryne, Atelopus, Dendrophryniscus,
Melanophryniscus, and Osornophryne, E in Oreophrynella, unknown in
Frostius); vocal slitpresent (absent in Dendrophryniscus, Oreophrynella, and
Osornophryne); a columella (absent in Atelopus [except flavescens group],
Dendrophryniscus, Melanophryniscus, Oreophrynella, and Osornophryne);
frontoparietal fontanelle obscured by medial articulation of frontoparietals
(no medial articulation in Oreophrynella); otic ramus of squamosal extending
over prootic (not so in Oreophrynella, unknown in Andinophryne); 8 presac-
ral vertebrae (normally 7 in Atelopus, 6 in Osornophryne, and 5 in Oreophry-
nella); no omosternum (present in Andinophryne); cervical not fused to
second presacral vertebra (fused in Atelopus and Oreophrynella); sacrum not
fused to coccyx (fused in Dendrophryniscus, Oreophrynella, and Osorno-

A NEW GENUS AND SPECIES OF HONDURAN TOAD 11

phryne); 2 centralia in hand (1 in Atelopus, Dendrophryniscus, Frostius, and
Osornophryne, unknown in Andinophryne); fingers free of webbing (web-
bing well -developed in Andinophryne and Osornophryne); gastromyzophor-
ous larvae (generalized oral disc in Dendrophryniscus, Frostius, and Melano-
phryniscus, unknown in Andinophryne, Oreophrynella, and Osornophryne).

Atelophryniscus can be distinguished from the "bufonines" (Bufo will be
discussed separately) by the following combination of characters: tympanic
annulus concealed by skin (tympanum absent in Crepidophryne and Rham-
phophryne, present and visible in Peltophryne); adductor longus absent
(present in Peltophryne); adductor mandibulae in S+E condition (E in Crepi-
dophryne, unknown inRhamphophryne); a columella (absent in Crepidophryne
andRhamphophryne); a well-developed palatine (reduced in Crepidophryne);
anormal nasal region (extensively ossified and protruding inRhamphophryne);
no paired rostral bones (paired in Peltophryne [except P. fluviatica]); 8
presacral vertebrae (7 in some species of Rhamp hop hryne); coccyx without
expanded lateral edges (present in Crepidophryne); pha-langeal formula of
manus 2-2-3-3, that of pes 2-2-3-4-3 (1-2-3-2 and 1-2-2-4-2, respectively, in
Crepidophryne); tips of distal phalanges of manus modified T-shaped (simple
or only slightly knobbed in Crepidophryne, Peltophryne, andRhamphophryne);
gastromyzophorous larvae (generalized oral disc in Peltophryne, unknown in
Crepidophryne and Rhamphophryne).

The tadpoles of Atelophryniscus resemble those described for various
species of Atelopus (Duellman and Lynch, 1969; Gray and Cannatella, 1985;
Lescure, 1981; Lynch, 1986; Mebs, 1980; Starrett, 1967). Tadpoles of both
genera have depressed bodies with flattened venters and an enlarged ventral
mouth followed by a large suctorial disc with a raised rim extending 2/3 to
4/5 of the length of the body. They also have the typical bufonid condition of
two upper and three lower denticle rows, a medial anal tube, and a sinistral
spiracle. Color in life also is similar among the tadpoles of Atelophryniscus
and those of some species of Atelopus. Duellman and Lynch (1969: 234)
reported A. certus as "black with golden bronze flecks" and Gray and
Cannatella (1985: 916) recorded "black with pale golden marks on dorsum of
body and caudal musculature" for A. peruensis. Both approach the black body
with gold markings and black caudal musculature with pale yellow bands and
spots found in Atelophryniscus. Tadpoles of Atelopus subornatus and other
species in Andean Columbia are boldly marked "cream and black" like those
of Atelophryniscus (Lynch, 1986: 129). Differences between the two genera
are in arrangement of the papillae (median half of upper and median third of
lower lip bare in Atelophryniscus, complete on upper lip and a broad bare area
on lower lip in Atelopus [Mebs, 1980, reported a complete row of papillae on
lower lip in A. cruciger; Lescure, 1981, discredited this statement]) and the
lateral folds of the mouth (well-developed in Atelophryniscus, absent in
Atelopus). Differences also exist in the tadpole sizes. Of the tadpoles of

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Atelopus species described, only that of A. subornatus is similar in size to that
of Atelophryniscus (Lynch, 1986); the remaining species are considerably
smaller. For example, the Atelopus certus tadpole in stage 36 has a mean body
length of 6.8 mm and a mean total length of 14.0 mm and A. pulcher in stage
37 has a body length of 6.9 mm and a total length of 13.7 mm (Duellman and
Lynch, 1969). A tadpole of Atelophryniscus in stage 37 has a body length of
9.6 mm and a total length of 26.4 mm.

Old World Genera. Frost (1985) recognized ten genera of bufonids in
Africa and six in Asia, exclusive of the genus Bufo. Dubois (1987), using the
features of reproduction and development (data taken from Grandison, 1978;
Wake, 1980), divided the African genus Nectophrynoides into four genera by
naming three new genera to accommodate the eight species previously placed
in this genus (Frost, 1985). For convenience, these eight species are retained
in the genus Nectophrynoides, in the following discussion. The African
bufonids (exclusive of Bufo) were divided into two assemblages by Gran-
dison (1981). The "Nectophryne line" contains the genera Capensibufo,
Didynamipus, Laurentophryne, Nectophryne, Nectophrynoides, Werneria,
and Wolterstorffina, whereas Mertensophryne, Stephopaedes, and possibly
Schismaderma were thought to be related to the "Bufo vertebralis group." No
study comparable to those of Grandison (1978, 1980a, 1981) exists for the
African genera Mertensophryne, Schismaderma, and Stephopaedes or for the
Asian genera Ansonia, Bufoides, Leptophryne, Pedostibes, Pelophryne, and
Pseudobufo.

Stephopaedes was diagnosed on the basis of having larvae with a unique
"crown" on the head (Channing, 1978). However, Grandison (1980b) dem-
onstrated that the larvae of Mertensophryne micranotis have a similar
"crown." The genera Bufoides and Schismaderma are also poorly defined;
thus, the data in the following discussion are incomplete for these genera.

Atelophryniscus can be distinguished from the Old World bufonids (Bufo
will be discussed separately) in having cranial crests (no cranial crests in all
genera, except Bufoides)', parotoid glands (no parotoid glands in Didynami-
pus, Laurentophryne, Nectophryne, Werneria, Wolterstorffina, Ansonia,
Bufoides, Pelophryne, and Pseudobufo, feebly developed in Mertensophryne
and Schismaderma, a tendency toward a loss in Nectophrynoides); no
adductor longus (present in Capensibufo and Leptophryne); a short tensor
fasciae latae (elongate in Didynamipus, Laurentophryne, Nectophryne,
Werneria, Wolterstorffina, Leptophryne, Pelophryne, and Pseudobufo); a
columella (absent in Didynamipus, Laurentophryne, Nectophryne, Werneria,
Wolterstorffina, Mertensophryne, and Stephopaedes, a tendency toward a
loss in Nectophrynoides); quadratojugal well-developed and in bony contact
with maxilla (reduced or vestigial in Laurentophryne, Nectophryne, Wolter-
storffina, Ansonia, Leptophryne, Pelophryne, and Pseudobufo, absent in
Didynamipus); sl well-developed palatine (reduced in Capensibufo, Didy-

A NEW GENUS AND SPECIES OF HONDURAN TOAD 13

namipus, and Laurentophryne, absent in Nectophryne and Pelophryne)',
frontoparietal fontanelle obscured by medial articulation of frontoparietals
(no medial articulation in Capen.sibu.fo and Nectophryne); 8 presacral verte-
brae (7 in Didy namipus, Laurentophryne, Mertensophryne , and Pelophryne,
7 or 8 in Nectophryne and Nectophrynoides); cervical not fused to second
vertebra (fused in Leptophryne); fully arciferal pectoral girdle (modified in
Didynamipus and Leptophryne); no omosternum (present in Capensibufo,
Nectophrynoides, and Werneria); sacral-coccygeal articulation bicondylar
(fused in Didynamipus, Laurentophryne, Nectophryne, and Pelophryne,
monocondy lar or fused in Wolterstorffina, monocondylar in Mertensophryne) ;
condyles of vertebrae ossified (not ossified in Pseudobufo); coccyx without
expanded lateral edges (present in Didynamipus and Pelophryne); normal
phalangeal formula of 2-2-3-3 and 2-2-3-4-3 (reduced in Didynamipus and
Mertensophryne); terminal phalanges of hands expanded, moderately T-
shaped (simple or knobbed in Capensibufo, Didynamipus, Mertensophryne,
Werneria, and Stephopaedes); tips of digits not dilated into discs (dilated into
discs in Pedostibes); non-aquatic adults (aquatic with fully-webbed feet in
Pseudobufo); gastromyzophorous larvae (generalized oral disc in Capen-
sibufo, Schismaderma, Bufoides, Leptophryne, and Pedostibes, vestigial in
Pelophryne, funnel-shaped in Nectophryne and Wolterstorffina, an expanded
oral disc not followed by a suctorial disc with a raised rim in Werneria and An-
sonia, a "crown" on head in Stephopaedes and Mertensophryne, either
aquatic with generalized oral disc, terrestrial non-feeders, or omitted in
Nectophrynoides, unknown in Didynamipus, Laurentophryne, and Pseu-
dobufo).

The Genus Bufo. As presently understood, there are 205 species in the
genus Bufo, 92 of which occur in the New World (Frost, 1985; Hoogmoed,
1985). Atelophryniscus can be distinguished from Bufo by possession of
modified T-shaped distal phalanges on the hand (simple or slightly knobbed
inBufo [Laurent, 1986;Tihen, 1960, 1962]) and gastromyzophorous tadpoles
(larvae with generalized oral disc and no abdominal disc in Bufo). Atelo-
phryniscus also lacks an adductor longus muscle, which is present in most
Bufo. This muscle can be absent, however, in some species (Limeses, 1964;
McDiarmid, 1971).

DISCUSSION

Cannatella (1986: 202), as noted above, separated the New World bufonid
genera into two subgroups. He did so with reservations, however, treating the
"atelopodine" genera "as a monophyleuc unit, primarily to point out areas
where further work is needed." That his reservations were warranted is further
indicated by the recognition of Andinophryne and Atelophryniscus. Andino-
phryne has partial fusion of the epicoracoid cartilages and lacks cranial crests

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

(Hoogmoed, 1985), both "atelopodine" features, but has parotoid glands, a
"bufonine" feature. Atelophryniscus shares several "bufonine" traits but has
a gastromyzophorous tadpole, a characteristic shared with Atelopus. The
naturalness of the Cannatella arrangement is further compromised by confu-
sion concerning the placement of Rhamphophryne. Cannatella (1986: 201)
associated this genus with Crepidophryne and Peltophryne on the basis of
sharing "fully arciferal pectoral girdles and the presence of cranial crests and
parotoid glands. . . ." Hoogmoed (1985: 258), however, scored the girdle
condition in Rhamphophryne as Ap following the terminology of McDiarmid
(1971: 45), who described this condition as "anteriorly firmisternal, posteri-
orly arciferal." Hoogmoed 's decision was apparently based on Trueb's (1971:
16) description of Rhamphophryne, which stated: "The pectoral girdle is
arciferal but tends to have a broad epicoracoid bridge." As such, Hoogmoed
(1985) associated Rhamphophryne with the "atelopodines" in spite of its
possession of cranial crests and parotoid glands. The monophyletic nature of
this genus also has been questioned by Cannatella (1 986: 200), who noted that
the "extensively ossified and protruding nasal region is the only unique
derived feature of the genus."

Another complication in assessing bufonid generic relationships is the
lack of understanding about the huge and undoubtedly polyphyletic Bufo.
Although usually considered to be characterized as having an adductor longus
muscle (e.g., Hoogmoed, 1985: 258; Laurent, 1986: 702; Tihen, 1960: 232),
one New World species, B. ockendeni (= B. veraguensis) is known to lack it
(Limeses, 1964). This species also lacks a tympanum (Savage, 1969). B.
veraguensis was placed in the same group with B.fissipes and B. quechua by
Gallardo, 1961: 5), who noted that the tympanum is "not visible" in these two
taxa. Three other species placed in this group by Gallardo (1961 , 1967) have
"visible" tympana. Duellman and Toft (1979: 62) described a new species, B.
nesiotes, indicating that it "presumably" belongs to the veraguensis group.
Bufo nesiotes has a "large, distinct" tympanum (although their drawing of the
lateral view of the head of the holotype does not match the description), unlike
some of the other members of this group. Unlike the "typical" New World
Bufo, B. nesiotes and B.fissipes lack cranial crests. All members of the
veraguensis group occur in montane habitats in Peru and Bolivia. Six Central
American and southern Mexican montane species (B. bocourti, B. fastidi-
osus, B. holdridgei, B. periglenes, B. peripatetes, and B. tacanensis) and two
South American species (B. cophotis andfi. variegatus) lack both a tympa-
num and a columella (Boulenger, 1900; Martin, 1972; Savage, 1972; Smith,
1952; pers. observ.). Some individuals of the Mexican B. occidentalis have
the tympanum concealed by skin (Duellman, 1961). Unlike other Bufo,
members of the B. haematiticus group have an omosternum (Trueb, 1971).
Finally, Bufo nesiotes has few, large, unpigmented eggs which, as noted by
Duellman and Toft (1979: 63), makes it "an unlikely candidate for the usual

A NEW GENUS AND SPECIES OF HONDURAN TOAD 15

mode of reproduction in Bufo — strings of small pigmented eggs in water."
They further indicated that "it is possible that 5. nesiotes lays terrestrial eggs
undergoing direct development."

Absence of data regarding aspects of reproduction and/or reproductive
behavior for many New World genera (including many of the montane species
of "Bufo" discussed above) makes it difficult to assess the phylogenetic
relevance of these features of bufonid biology. Finally, the scant effort to
search for the relationships between New and Old World genera of bufonids
renders suspect any arrangement based only on genera from one hemisphere
or the other. Further discussion of the problems presented by our poor state
of knowledge of New World bufonids is in Hoogmoed (1985: 261-262).

Given the existence of these problems, it is futile at this juncture to attempt
a phylogenetic analysis of relationships within any generic segment of the
family Bufonidae. We limit our comments on the relationships of Atelo-
phryniscus, therefore, to consideration of its probable closest relative. Ate lo-
phryniscus appears to be a derivative of some group of Bufo, agreeing with
this taxon in possessing a number of primitive features (including, in addition
to the "bufonine" features [we consider the presence of parotoid glands
primitive, following Grandison, 1981, not derived as suggested by Can-
natella, 1986] a short tensor fasciae latae, S+E adductor mandibulae, 8
presacral vertebrae, palatine bone well-developed, 2-2-3-3 phalangeal for-
mula, coccyx without expanded lateral edges, 2 centralia in hand). The
principal distinction between these two genera is in the possession of a
gastromyzophorous tadpole by Atelophryniscus. The resemblance between
this tadpole and those olAtelopus might be considered as an argument for a
relationship between this genus and Atelophryniscus. We consider the resem-
blance to represent convergence resulting from adaptation to similar environ-
ments. Mountain-stream tadpoles typically exhibit adaptation for life in
rapidly- flowing streams by having a depressed body, muscular tail with low
fins, and either suctorial mouthparts or a suctorial abdominal disc. Suctorial
mouthparts are known in a variety of mountain-stream-breeding anurans. As
noted above, only a few species of anurans are known to possess the
gastromyzophorous condition. We thus consider the resemblance of Atelo-
phryniscus and Atelopus to result from the superim position of stream-adapted
features (including the suctorial abdominal disc) on the standard bufonid
tadpole morphology (2/3 tooth rows, median anal tube, and sinistral spiracle).

ACKNOWLEDGMENTS

Gustavo A. Cruz, Mario R. Espinal, Kelly M. Hogan, and Walter Holmes
provided field assistance. Gustavo A. Cruz and Wilfredo Aguilar of the
Direction General de Recursos Naturales Renovables provided collecting
permits. Jay M. Savage and Brian I. Crother of the University of Miami and

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Arnold G. Kluge and Dennis M. Harris of the University of Michigan
Museum of Zoology loaned comparative material. Jay M. Savage assisted
with the identification of the condition of the pectoral girdle and adductor
longus muscle in Atelophryniscus. David C. Cannatella and John D. Lynch
provided unpublished data on several of the New World bufonid genera.
David C. Cannatella also provided us with an advance copy of a manuscript
and copies of several pertinent published papers. Ronald Altig and Peter Gray
provided information on bufonid tadpoles. Two anonymous reviewers con-
tributed many helpful suggestions.

LITERATURE CITED

Boulenger, G. A. 1900. Descriptions of new batrachians and reptiles collected by Mr. P. O.
Simons in Peru. Ann. Mag. Nat. Hist. (7)6: 181-186.

Cannatella, D. C. 1986. Anew genus of bufonid (Anura) from South America, and phylogenetic
relationships of the Neotropical genera. Herpetologica 42: 197-205.

Charming, A. 1 978. A new bufonid genus (Amphibia: Anura) from Rhodesia. Herpetologica 34:

394-397.
Dubois, A. 1987. Miscellanea taxinomica [sic] batrachologica (I). Alytes 5: 7-95 (1986).

Duellman, W. E. 1961 . The amphibians and reptiles of Michoacan, Mexico. Univ. Kansas Publ.
Mus. Nat. Hist. 15: 1-148.

. 1970. The hylid frogs of Middle America. Mus. Nat. Hist. Univ. Kansas Monogr. (1):

i-xii, 1-753.

Duellman, W. E., and J. D. Lynch. 1969. Descriptions of Atelopus tadpoles and their relevance
to atelopodid classification. Herpetologica 25: 231-240.

Duellman, W. E., and C. A. Toft. 1979. Anurans from Serrania de Sira, Amazonian Peru:
Taxonomy and biogeography. Herpetologica 35: 60-70.

Frost, D. R. (ed.). 1985. Amphibian species of the world. A taxonomic and geographical
reference. Allen Press, Inc. and Association of Systematic Collections, Lawrence, Kansas.

Gallardo, J. M. 1 961 . Three new toads from South America: Bufo manicorensis, Bufo spinulosus
altiperuvianus and Bufo quechua. Breviora (141): 1-8.

. 1967. Bufo gnustae sp. n. del grupo de B. ockendeni Boulenger, hallado en la Prov. de

Jujuy, Argentina. Neotropica 13: 54-56.

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on
identification. Herpetologica 16: 183-190.

Grandison, A. G. C. 1978. The occurrence of Nectophrynoides (Anura Bufonidae) in Ethiopia.
A new concept of the genus with a description of a new species. Monitore Zoologico Italiano
(n. s. )Suppl. 11:119-172.

. 1980a. Anew genus of toad (Anura: Bufonidae) from the Republic of South Africa with

remarks on its relationships. Bull. British Mus. (Nat. Hist.) 39: 293-298.

. 1980b. Aspects of breeding morphology in Mertensophryne micranotis (Anura:

Bufonidae): secondary sexual characters, eggs and tadpole. Bull. British Mus. (Nat. Hist.)
39: 299-304.

. 1981. Morphology and phylogenetic position of the West African Dldynamipus

sjoestedti Andersson, 1903 (Anura Bufonidae). Monitore Zoologico Italiano (n.s.) Suppl.
15: 187-215.

A NEW GENUS AND SPECIES OF HONDURAN TOAD 17

Gray, P., and D. C. Cannatella. 1985. Anew species oi Atelopus (Anura, Bufonidae) from the
Andes of northern Peru. Copeia 1985: 910-917.

Holdridge, L. R. 1967. Life zone ecology. Tropical Science Center, San Jose, Costa Rica.

Hoogmoed, M. S. 1985. A new genus of toads (Amphibia: Anura: Bufonidae) from the Pacific
slopes of the Andes in northern Ecuador and southern Colombia, with the description of two
new species. Zoologische Mededelingen 59: 251-274.

Inger, R. F. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana:
Zool. 52: 1-402.

Kuramoto, M., C. Wang, and H. Yti. 1984. Breeding, larval morphology and experimental
hybridization of Taiwanese brown frogs, Rana longicrus and R. sauteri. J. Herpetol. 18:
387-395.

Laurent, R. F. 1986. Sous-classe des Lissamphibiens (Lissamphibia). Systematique. Pp.
594-797. In Grasse, P. and M. Delsol (eds.). Traite de Zoologie. XIV. Amphibiens. Fas. IB.
Masson, Paris.

Lescure, J. 1981. Contribution a l'etude des amphibiens de Guyane francaise IX. Le tetard
gastromyzophore d' Atelopus flavescens Dumeril et Bibron (Anura, Bufonidae). Amphibia-
Reptilia 2: 209-215.

Limeses, C. E. 1964. La musculature del muslo en los ceratofrinidos y formas afines con un
analisis critico sobre la signification de los caracteres miologicos en la sistematica de los
anuros superiores. Univ. Buenos Aires, Fac. Cienc. Exactas Natur., Contrib. Cient., Ser.
Zool. 1: 193-245.

Liu, C. 1950. Amphibians of western China. Fieldiana: Zool. Mem. 2: 1-400.

Lynch, J. D. 1971. Evolutionary relationships, osteology, and zoogeography of leptodactyloid
frogs. Univ. Kansas Mus. Nat. Hist. Misc. Publ. (53): 1-238.

. 1986. Notes on the reproductive biology of Atelopus subornatus. J. Herpetol. 20:

126-129.

Martin, W. F. 1972. Evolution of vocalization in the genus Bufo. Pp. 279-309. In Blair, W. F.
(ed.). Evolution in the genus Bufo. Univ. Texas Press, Austin.

McDiarmid, R. W. 1971. Comparative morphology and evolution of frogs of the Neotropical
genera Atelopus, Dendrophryniscus \ Melanophryniscus, and Oreophrynella. Bull. Los
Angeles Co. Mus. Nat. Hist. Sci. (12): 1-66.

Mebs, D. 1980. Zur Fortpflanzung von Atelopus cruciger. Salamandra 16: 65-81.

Myers, C. W., and W. E. Duellman. 1 982. A new species oiHyla from Cerro Colorado, and other
tree frog records and geographical notes from western Panama. Amer. Mus. Novitates
(2752): 1-32.

Pillai, R. S., and G. M. Yazdani. 1973. Bufoides, a new genus for the rock-toad, Ansonia
meghalayana Yazdani & Chanda, with notes on its ecology and breeding habits. J. Zool. Soc.
India 25: 65-70.

Pregill, G. 1981. Cranial morphology and the evolution of West Indian toads (Salientia:
Bufonidae): resurrection of the genus Peltophryne Fitzinger. Copeia 1981: 273-285.

Ruiz-C, P. M., and J. I., Hemandez-C. 1976. Osornophryne, genero nuevo de anfibios
bufonidos de Colombia y Ecuador. Caldasia 11(54): 93-148.

Savage, J. M. 1969. Clarification of the status of the toad, Bufo veraguensis O. Schmidt, 1857.
Copeia 1969: 178-179.

. 1972. The systematic status of Bufo simus O. Schmidt with description of a new toad

from western Panama. J. Herpetol. 6: 25-33.

Savage, J. M., and A. G. Kluge. 1961. Rediscovery of the strange Costa Rica toad, Crepidius
epioticus Cope. Rev. Biol. Trop. 9: 39-51.

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Smith, P. W. 1952. A new toad from the highlands of Guatemala and Chiapas. Copeia 1952:

175-177.
Starrett, P. 1967. Observations on the life history of frogs of the family Atelopodidae.

Herpetologica 23: 195-204.
. 1968. The phylogenetic significance of the jaw musculature in anuran amphibians.

Ph.D. Diss. Univ. Michigan, Ann Arbor.
Tihen, J. A. 1 960. Two new genera of African bufonids, with remarks on the phylogeny of related

genera. Copeia 1960: 225-233.

. 1962. Osteological observations on New World Bufo. Amer. Midland Nat. 67: 157-183.

Trueb, L. 1971. Phylogenetic relationships of certain Neotropical toads with the description of

a new genus (Anura: Bufonidae). Los Angeles Co. Mus. Contrib. Sci. 216: 1^40.

. 1973. Bones, frogs, and evolution. Pp. 65-132. In Vial, J. L. (ed.). Evolutionary biology

of the anurans. Contemporary research on major problems. Univ. Missouri Press, Columbia.

Wake, M. H. 1980. The reproductive biology of Nectophrynoides malcomi (Amphibia:
Bufonidae), with comments on the evolution of reproductive modes in the genus Necto-
phrynoides. Copeia 1980: 193-209.

Wassersug, R. J., and W. R. Heyer. 1983. Morphological correlates of subaerial existence in
leptodactylid tadpoles associated with flowing water. Canadian J. Zool. 61: 761-769.

University of Kansas Publications

MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural
History, beginning with volume 1 in 1946, was discontinued with
volume 20 in 1971. Shorter research papers formerly published in
the above series are now published as Occasional Papers, Museum
of Natural History. The Miscellaneous Publications, Museum of
Natural History, began with number 1 in 1946. Longer research
papers are published in that series. Monographs of the Museum of
Natural History was initiated in 1970. Authors should contact the
managing editor regarding style and submission procedures before
manuscript submission. All manuscripts are subject to critical
review by intra- and extramural specialists; final acceptance is at
the discretion of the Director. The Occasional Papers and Miscel-
laneous Publications are produced by a Macintosh computer, Las-
erWriter, and the software Microsoft® Word and PageMaker®.
Conversion to current format by E. O. Wiley.

Institutional libraries interested in exchanging publications
may obtain the Occasional Papers and Miscellaneous Publications
by addressing the Exchange Librarian, University of Kansas Li-
brary, Lawrence, Kansas 66045. Individuals may purchase sepa-
rate numbers of all series. Prices may be obtained upon request ad-
dressed to Publications Secretary, Museum of Natural History,
University of Kansas, Lawrence, Kansas 66045 - 2454 U.S.A.

Editor. Robert D. Holt

Managing Editor. Joseph T. Collins

Design and Typesetting: K. A. Shaw and Joseph T. Collins

PRINTED BY

UNIVERSITY OF KANSAS PRINTING SERVICE

LAWRENCE. KANSAS

DATE DUE

-MAR 34-1996

DEMCO. INC 38-2931

3 2044 093 361 707