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Source : MNHN, Paris

AINTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

November 1996 Volume 14, N° 3

Alytes, 1996, 14 (3): 101-114. ee

liothèque Centrale Muséum

3 3001 00111580

Taxonomy and geographic distribution

of a northwestern Venezuelan toad

(Anura, Bufonidae, Bufo sternosignatus)

Enrique LA MaRCA * & Abraham MUHARES-URRUTIA **

* Laboratorio de Biogeografñia, Escuela de Geografia, Facultad de Ciencias Forestales,

Universidad de Los Andes, Apartado 116, Mérida 5101-A, Venezuela

** Centro de Investigaciones en Ecologia y Zonas Aridas (CIEZA),

Universidad Francisco de Miranda, Apartado Postal 7506, Coro 4101-A, Venezuela

In this paper we provide a redescription of Bufo sternosignatus, a

northwestem Venezuelan toad usually associated with the “Bufo typhonius

complex”, and detailed data on its extemal morphology, skin, meristic

characters, pattern of coloration in life and after preservation, and biogeogra-

phy.

INTRODUCTION

Bufo sternosignatus was described by Albert GÜNTHER (1859) in a book dealing with

the anurans deposited in the collection of the British Museum of Natural History

(London). The type series included a male and a female from “Venezuela”, a female from

“Puerto Cabello” (Estado Carabobo, Venezuela), and two Mexican juvenile toads stated

as coming from “Mexico” and “Cordova”. The latter were considered by BOULENGER

(1882: 320) as Bufo valliceps Wiegmann, 1833, a decision followed by KELLOGG (1932: 68).

The treatment of Bufo sternosignatus as a junior synonym of Bufo valliceps by PORTER

(1964: 242) probably reflects this partial assignment, inasmuch as BOULENGER (1882)

restricted the name to the Venezuelan types, although PORTER (1964) did not do this.

BOULENGER (1882: 323) also assigned to B. sternosignatus two specimens coming from

A Source : MNHN, Paris

102 ALYTES 14 (3)

Fig. 1. — Bufo sternosignatus, ULABG 1924, adult female, SVL 48.6 mm.

“Bogotä” (Colombia). Two other records from Colombia, Cafetal Amelia, near Angelé-

polis, 1820 m, by PERACCA (1914), and Cañôn de Tolima, by WERNER (1916), were denied

by RIVERO (1961: 29), who considered them as probably referring to Bufo typhonius alatus.

CocHRAN & Goin (1970: 112) retained the Bogotä frog in the Colombian fauna, “on the

strength of BOULENGER‘"S identification”.

The specific status of Bufo sternosignatus was apparently not questioned until SHREVE

(1947) treated it as a subspecies of Bufo typhonius, with the provision that “even though

Bufo typhonius alatus Thominot, 1884 from Panama should prove to be synonymous,

sternosignatus, being the older name, will have to be used”. RIVERO (1961: 28-29)

considered the species to be very similar to Bufo typhonius alatus, adding that it “perhaps

interbreeds (though not freely) in part or all of its range”, and later (RIVERO, 1964b: 314)

that they “‘seemed to be most closely related”. All other references to the species in the

literature have appeared in catalogues or locality lists under Bufo sternosignatus (e.g. LUTZ,

1927a-b, 1955; PARKER, 1935; SHREVE, 1947; SmiTH & TAYLOR, 1950; GINÉS, 1959; RIVERO,

1964a-b, 1967; SOLANO, 1969; MÜLLER, 1973; LyNCH, 1979; HARDING, 1983; FROST, 1985;

La MaRCA, 1992a, 1995).

Bufo sternosignatus was placed in the Bufo typhonius group by Cet (1972) and as such

it is still recognized. The most recent grouping of South American bufonids (by D'UELLMAN

Source : MNHN, Paris

La MarcA & MuARES-URRUTIA 103

& SCHÜLTE, 1992) placed the species in the typhonius group along with the following

species: B. ceratophrys Boulenger, 1882; B. dapsilis Myers & Carvalho, 1945; B. iserni

(Jiménez de La Espada, 1875); B. nasicus Werner, 1903; B. roqueanus Melin, 1941; and B.

typhonius (Linnaeus, 1758). HoOGMOED (1989) was of the opinion that even those species

do not form a natural group and recommended that the name Bufo typhonius be

suppressed, because of nomenclatural problems and the confusion surrounding this name.

Regarding B. sternosignatus, Cet (1972) had otherwise indicated that “it might alterna-

tively be related to B. granulosus”. The systematic status of B. sternosignatus remains

problematic, as was pointed out by HOOGMOED (1990), who also indicated that “‘statements

about relationship seem to have been made on the basis of general appearance (colour and

pattern) and not on any hard morphological data”. After examining the Venezuelan

syntypes, HooGMoED (1990) concluded that these certainly were not Bufo valliceps, that

they were not related to Bufo granulosus either, and did not rule out the possibility that

the species is identical to B. “typhonius” from Panama and adjacent North America.

Although relatively easy to set apart from other Venezuelan bufonids, Bufo

sternosignatus (fig. 1) has not been properly studied, probably due to the paucity of

specimens in collections. We had the opportunity to collect the species in the field, and

from the examination of these and other museum specimens we conclude that, although

it shares the pale vertebral line and a so-called “‘dead-leaf pattern” with some species in

the “typhonius group”, it is a perfectly valid and distinct taxon probably not related to any

species in this complex. We do not rule out the possibility that it might be related to Bufo

granulosus. Bufo sternosignatus is readily distinguished from other northern South

American bufonids by being more spinose, by having very distinct tympana, oval-shaped

spinose parotoid glands, a pale cross design on ventral surfaces, and by the tubercles on

flanks disposed in lateral rows running somewhat parallel to the dorsum. Although the

type locality of Bufo sternosignatus was restricted by COCHRAN & Go (1970: 111) to

“Puerto Cabello, Venezuela”, which “almost” amounts to a lectotype designation, no

proper such designation has ever been published for this species. Unfortunately, we could

not study the syntypes, nor could we designate a lectotype, an action we feel needs to be

done in the frame of a revisionary work on the systematic relationships of the species, a

task beyond the scope of this paper. Nonetheless, we offer here a redescription and

additional data on the species, in the hope that they may help for such a revisionary work.

MATERIAL AND METHODS

Museum abbreviations employed in the text are as follows: CIEZAH, Coleccién

Herpetolôgica del Centro de Investigaciones en Ecologia y Zonas Aridas, Universidad

Francisco de Miranda, Venezuela; CVULA, Colecciôn de Vertebrados, Universidad de

Los Andes, Venezuela; MCNG, Museo de Ciencias Naturales Guanare, Venezuela;

ULABG, Coleccién de Anfibios y Reptiles, Instituto de Geografia, Universidad de Los

Andes, Venezuela; UMMZ, University of Michigan Museum of Zoology, USA.

The following specimens of Bufo sternosignatus, all new records from Venezuela, and

upon which is based the redescription of the species, were examined during the course of

this study:

Source : MNHN, Paris

104 ALYTES 14 (3)

Estado Aragua: ULABG 3947, La Trilla, near Ocumare de La Costa, 150 m. Estado

Barinas: CVULA 5158, Barinitas. Estado Carabobo: ULABG 1319, Hacienda Bucarito,

near Montalbän, 700 m. Estado Falcon: CIEZAH 322, road to Cerro Galicia, near

Curimagua, 1150 m; MCNG 2190-2191, Finca La Felipina, approx. 3 km from La

Soledad de Uria, Sierra de San Luis, ca. 1300 m; CIEZAH 323, CVULA 5118-5119,

Cataratas de El Hueque, Parque Nacional J. C. Falcon, Sierra de San Luis, 800 m;

CIEZAH 317-319, San Diego, Sierra de San Luis, approx. 1100 m; ULABG 2940, Cerro

Chichiriviche. Estado Portuguesa: ULABG 3726-3727, 6 km from Rio Saguaz, on road

Biscucuy-Chabasquén, 585 m. Estado Trujillo: ULABG 1924, carretera Bocon6-Trujillo,

near intersection with road Trujillo-La Concepcion; ULABG 2667, Agua Negra near

Sabaneta, 1405 m, on road from Trujillo to San Läzaro.

Terminology, measurements and web formula system follow LA MARCA (1984, 1989,

1994b).

REDESCRIPTION OF BUFO STERNOSIGNATUS GÜNTHER, 1859

DIAGNOSIS AND DEFINITION

A moderate-sized species of Bufo (mean SVL: 5 males, 49.5 mm; 5 females, 42.3 mm)

having the following combination of characters: (1) distinct supraorbital and anteorbital

crests bordering upper eyelid and anterior part of eye; (2) supratympanic crest prominent;

(3) parotoid glands oval-shaped and densely tuberculate; (4) snout truncated in dorsal

view; (5) tympanum large (about 2/3 the horizontal length of eye); (6) skin densely

tubercular with numerous minute spicules; (7) a pale-cream cross usually present on chest.

Bufo sternosignatus occurs sympatrically with B. marinus, but may be distinguished by

the following characters (in B. sternosignatus): smaller size, shorter head, smaller

tympanum, stronger supratympanic crest, snout truncated in dorsal view, and body

surfaces bearing minute spicules. B. sternosignatus more closely resembles B. typhonius

alatus, but differs by having a non-prominent snout, less prominent supraorbital crest,

weak occipital crest, parotoid glands oval-shaped, hands and feet with minute spicules,

metatarsal fold absent, and a pale-cream cross usually present on chest. B. sternosignatus

could also be confused with B. granulosus, but differs from the latter by having a larger

size, longer head, stronger supratympanic crest, smaller spicules on hand and feet, more

truncated snout, crests around nostrils absent, and no well-defined crest along canthus

rostralis.

DESCRIPTION

Head twice as long as wide and twice as wide as deep, narrower than body width; top

of head flat to slightly concave in the middle, shifting to slightly convex in the postoccipital

region, with prominent warts (ca. 0.5 mm in diameter), some of them coalescing to form

canthus rostralis. Canthus rostralis curved, well defined. Distinct supraorbital and

anteorbital crests bordering upper eyelid and anterior part of eye. Loreal region slightly

Source : MNHN, Paris

La MaRCcA & MUARES-URRUTIA 105

concave, descending almost vertically to upper lip; snout truncated in dorsal view, slightly

protruding (sensu HEYER et al., 1990: fig. 80) in lateral view; nares oval to rounded,

directed dorsolaterally; nostrils elevated, closer to tip of snout than to eye (about 1/5 of

distance from point of snout to eye); internarial distance about half interorbital distance.

Internarial region between moderate to highly concave. Interorbital distance slightly

(approx. 1/5) longer than upper eyelid width. Upper eyelid bearing pungent tubercles.

Upper eyelid with a thickened and finely granular border. External border of upper eyelid

with minute spicules, slightly projecting beyond eye. Lips not flaring. Eyes protuberant to

the sides and above dorsal surface of head. Eye diameter slightly longer than distance from

anterior border of eye to naris opening. Lower eyelid translucent (opaque in CIEZAH

323). Palpebral membrane dusted with brown, with a well defined dark brown border.

Pupil oval. Temporal region descending vertically. Supratympanic crest moderately to well

defined, fleshy, not projecting over temporal region, from posterior border of eye to

anterior or medial part of parotoid gland, with smooth borders, not covering tympanum.

Tympanum large (horizontal length about 2/3 of horizontal length of eye), conspicuous,

rounded to oval, bordered by tubercles, with well-defined elevated borders, and without

cartilaginous ring. Supratympanic crest prominent, short and thick, not projecting laterally

beyond most external border of upper eyelid, extending with a narrow crest descending

almost vertically, bordering tympanum anteriorly; supratympanic crest connected ante-

riorly with a crest that borders upper eyelids and anterior part of eye. Parotoids

oval-shaped, well visible from above, slightly swollen, about twice as long as wide, 1.5 to

2 times larger than width of upper eyelid.

Choanae rounded to oval, large (about 2/3 of internarial distance), located near the

margins of roof of mouth, covered or not by palatal shelf of maxillary arch, separated by

a distance larger than that between nostrils. Tongue oval, narrow (width about 1/3 of

length), with parallel margins; tongue entire in its anterior part, almost completely free

(more than 2/3 of its posterior part); teeth absent from maxilla, premaxilla and vomers.

Males with lateral vocal slits, at about 40 % of distance from base to tip of tongue, and

a possible subgular vocal sac (see below).

Cloacal opening located at or close to midlevel of thighs. Cloacal opening small,

directed posteroventrally, at about 2/3 of distance from base to top of thigh. Ulnar fold

absent. Hand (fig. 2) with outer metacarpal (palmar) tubercle well defined, large, not

divided, slightly heart-shaped; inner metacarpal (thenar) tubercle moderately to well

visible, small, about 1/4 to 1/2 size of palmar (sometimes inconspicuous), oval.

Supernumerary tubercles present, relatively rounded to squared, of small to very small

size. Subarticular tubercles double, rounded, small, conspicuous. Fingers without lateral

fringes; fingers almost completely free, except for a short and thick membrane between

fingers III and IV; hand web formula I 0 - O II 0 - O III 1 - 1 IV 0 - 0 V; membrane thick,

rugose, tuberculate. Tip of fingers rounded. Digital pads narrower than maximum width

of fingers. Relative length of fingers 1 > II < III > IV.

Foreleg with numerous small tubercles, some pungent. Tarsal fold absent; narrow

groove dividing tarsus longitudinally, from base of fingers IV and V to tibio-tarsal joint.

Foot (fig. 3) with an inner metatarsal tubercle prominent, flattened, smooth, oval, slightly

elongate in dorsal view, subconical in lateral view, 2.5 times larger than outer metatarsal

Source : MNHN, Paris

106 ALYTES 14 (3)

Fig. 2. — Left hand of Bufo sternosignatus, ULABG 1319. Note the numerous minute spicules on

skin. Line: 5 mm.

Source : MNHN, Paris

La MarCcA & MuARES-URRUTIA 107

Fig. 3. — Right foot of Bufo sternosignatus, ULABG 1319. Line: 5 mm.

Source : MNHN, Paris

108 ALYTES 14 (3)

tubercle, about twice as long as wide. Outer metatarsal tubercle twice as long as wide,

smaller than inner metatarsal tubercle, distinct, oval in dorsal view, prominent, flat and

smooth. Supernumerary tubercles present, small to medium-sized, with minute spicules.

Subarticular tubercles conspicuous, small and oval-shaped. Toe IV with a lateral

tuberculate fringe; toe tips rounded; thick tuberculate membrane between toes; maximum

web development reaching base of digital pads on left side of toes I, II and III, and right

side of V; fourth toe almost completely free; web formula I 1 - 2 II 2 - 2.5 III 2 - 2 IV

1.5 - 1 V; all toes bearing a lateral fringe-like row of tubercles that are a continuation of

those on free border of interdigital membrane. Heels not touching or only slightly so when

thighs are held at right angles to body axis; tibio-tarsal articulation reaching shoulders

when legs are adpressed forward (in the original description, heel is stated to reach eye;

in MCNG 2191, heel reaches tympanum; in CIEZAH 318, it reaches between tympanum

and shoulder). Relative length of toes: I < II < V < III < IV. Tibia about 37 % body

length.

Large cream eggs (up to 1.4 mm diameter) were found in convoluted oviducts of adult

females ULABG 1319 and 2406, and (up to 1.9 mm) in CIEZAH 319. Adult males have

elongate white testicles (e.g. CIEZAH 317-318), sometimes granulate (e.g. CIEZAH 323);

vocal sac is not evident, possibly subgular. Vocal slits may be present on one (e.g.

CIEZAH 323) or both sides (e.g. CIEZAH 318). Adult males CIEZAH 317-318 have

anterodorsal parts of fingers I and II partially keratinized.

Skin

Skin of dorsum, flanks, throat, venter, extremities (above and below), palms and soles

tubercular. Tubercles on dorsum and on top of head between rounded and pungent, from

0.5 to 1.5 mm in diameter, the largest on sides and posterior part of body; a stria

longitudinally dividing upper head and body in two equal halves, from snout tip to

urostyle or upper border of cloacal opening. Lower back and flank tubercles sometimes

ending in one or several (up to 8) spicules. Parotoid glands densely tuberculate. A row of

tubercles (sometimes inconspicuous), medium to large-sized, conical shaped, from

parotoids to groin; largest tubercles in middle of row. Loreal region with tubercles bearing

numerous minute spicules. Upper eyelid with numerous subconical tubercles; those on

external border ending in spicules. Temporal region and area near rictus covered by

numerous round tubercles, small to moderate-sized. Tympanum densely granular, granules

rounded, minute. Tubercles rounded and long (about 1.0 mm long) between upper part of

tympanum and supratympanic crest. Neck, in lateral view, with numerous, relatively

medium-sized, tubercles in its superior part; scarce and small in its inferior part. Flanks

with rounded tubercles, moderate-sized, some pungent.

Venter and throat with small (about 1.0 mm) tubercles, ending in numerous minute

spicules; cloacal opening smooth or bordered by rounded tubercles bearing numerous

spicules. Tubercles on extremities elevated, subconical, smaller than those on dorsum,

generally ending in a single spicule; lower aspect of forearm usually with large subconical

tubercles, arranged in two or three longitudinal rows; a longitudinal groove, slightly

oblique, from wrist to elbow; fingers totally spiculate, except on distal end, which is

smooth; interdigital membranes tuberculate; supernumerary and subarticular hand

Source : MNHN, Paris

La MarcA & MUARES-URRUTIA 109

tubercles bearing numerous minute pungent spicules. Innner metacarpal tubercle smooth

or occasionally with minute spicules. Free margin of hand-web with some minute acute

tubercles. Subarticular tubercles on feet with numerous minute spicules.

Measurements in mm (mean + one standard deviation)

Snout-to-vent length: adult males, 49.5 + 3.3 (N = 5, range 45.7-55.1); females, 42.3

+ 3.4(N = 5, range 37.1-47.6). Other measurements (N = 14, both sexes together): head

width, 15.1 + 2.0; head length, 12.4 + 1.5; eye, 4.5 + 0.6; eye to tip of snout distance,

5.5 + 0.6; eye to nostril distance, 3.8 + 0.6; internarial distance, 2.9 + 0.4; interorbital

distance, 5.2 + 0.8; eye to tympanum distance, 1.4 + 0.4; horizontal length of tympanum,

2.6 + 0.4; hand length, 9.9 + 1.3; foot length, 14.8 + 2.1; tibia length, 15.9 + 2.2.

Coloration

Patterns of dorsal and ventral coloration for the species have been illustrated by

GÜNTHER (1859: plate 5, fig. C) and LUTZ (19274: plate 8, figs. 3-4). RIVERO (1961: 28)

noted that this frog is usually of a reddish colour when alive, and that some specimens are

peculiarly marked with yellow and brown below. Data on coloration in life is available for

ULABG 1924 (fig. 1) and 3726-3727. Dorsum pale brown bearing a large irregular dark

brown to black marking bordered with a pale yellowish line; a cream line from point of

snout to end of urostyle; inferior part of flanks dark brown; forearms and thighs with

chocolate bands; throat and chest cream; a dirty-cream cross-shaped marking on dark gray

chest; upper part of flanks pale brown; lower parts of flanks chocolate. Tympanum

marbled with chocolate and dark brown. Numerous cream tubercles, on a plumbeus

background, on venter and under the thighs, and two irregular white spots on rear end of

upper lip, connected to a white blotch below eye (ULABG 1924). Golden iris, marbled

with brown in inferior half, parotoid glands ochre to pale brown, upper lip cream,

inconspicuously barred, and groin ochre with a pinkish hue (ULABG 3726-3727). Colour

slides of ULABG 1319 (fig. 4) show lower dorsum and cephalic irregular black spots, and

a butterfly-shaped dark brown to black marking between eyelids, all bordered with yellow

lines. Digits, proximal end of ventral surface of the thighs and axillary surfaces

ochre-reddish. Vertebral line, cross on ventral surfaces, lower venter, most parts of ventral

parts of thighs and shanks dirty white. Two white spots present at the rear end of upper

mandible.

Coloration of preserved specimens is detailed as follows. Dorsum from pale to dark

brown, usually with a pale vertebral line, sometimes bordered by black, from point of

snout to urostyle, sometimes hidden by a pale brown marking between the eyes. Some

specimens bearing dark-brown, bilaterally symmetrical or almost symmetrical markings,

on a pale brown background, these markings having sometimes a blackened border (e.g.

ULABG 1924); some specimens with dark-brown rounded markings, bordered by a pale

cream line at level of sacrum (e.g. CIEZAH 317, CVULA 1319). Some specimens (e.g.

CIEZAH 318) with pale brown tubercles surrounded by black, and some (e.g. CIEZAH

319 and 322, CVULA 5119, MCNG 2190-2191, ULABG 2940) with a dorsal pale brown

subtriangular marking, surrounded by black, with apex directed backwards. All specimens

Source : MNHN, Paris

110 ALYTES 14 (3)

Fig. 4. — Bufo sternosignatus, ULABG 1319, adult female, SVL 55.1 mm. Bottom: dorsal view (note

pale vertebral line and dark dorsal markings). Top: ventral view (note pale cross on chest).

Source : MNHN, Paris

La MarcA & MUuARES-URRUTIA 111

examined with dark brown symmetrical markings between the supratympanic crests and

the posterior part of the upper eyelids, that may have had a dark (e.g. ULABG 1924) or

pale brown (e.g. ULABG 1319) border. Two infraocular dark brown bands present,

separated by a paler band (sometimes cream-coloured, like CVULA 5119), although one

of these bands (or both) may be inconspicuous (e.g. CIEZAH 322). Palpebral membrane

dusted with brown, with a well defined dark brown border.

Upper part of flanks same as dorsum; lower part paler, similar to coloration of venter.

Extremities, above, generally with dark brown bars on pale brown background; in some

instances (e.g. CIEZAH 317-318), the bars are not well-defined. Ventral aspect of

hindlimbs immaculate cream. Ventral aspect of forelimbs longitudinally bicoloured

(internal half, cream; external half, brown); a dark brown line present from palmar

tubercle to almost the elbow. Palms dark-brown, but less than soles, which are the darkest

ventral surfaces. Lateral fringe on toe IV made up of tubercles paler than those on top of

digit.

Throat and chest usually marbled with brown; border of lower mandible inconspic-

uously marbled. Anterior part of chest and gular region somewhat darker in CIEZAH 317.

Venter usually marbled (most specimens bearing dispersed pale brown specks on a cream

background) to completely pale gray (e.g. CIEZAH 318-319). Lower venter, between groin

and cloacal region, immaculate cream. A pale-cream cross usually present on chest

(conspicuous in CIEZAH 319 and 322 and in CVULA 1319; inconspicuous in CIEZAH

317, ULABG 1924 and CIEZAH 323). The longitudinal line of this pale-cream cross

extends from base of throat to middle of the venter; the transversal line connects insertions

of arms.

BIOGEOGRAPHY

Confusion of Bufo sternosignatus with other taxa brought misconceptions about the

distribution of the species, that at times has been thought to occur in Central America and

Colombia (e.g. BOULENGER, 1882), Mexico (LuTz, 1927a), and even in “the Guianas,

rarely extending farther southwards in eastern Brazil” (PARKER, 1935: 528). Central

American records probably belong to Bufo valliceps and Bufo typhonius alatus. Colombian,

Guianan and Brazilian records probably correspond to some other forms in the “typhonius

complex”. Venezuelan records in the literature for Bufo sternosignatus include mostly

specimens from the Cordillera de La Costa: thus, those from Estado Carabobo include

Puerto Cabello (GÜNTHER, 1859; BOULENGER, 1882), Rio La Mona and San Esteban (“Rio

Loma” and “San Esteban trail to Las Cuiguas”, respectively, according to UMMZ lists),

between Valencia and Caserio Silva (“Palma Sola” in UMMZ lists) and Rio Bejuma

(RIVERO, 1961). From Estado Aragua they include Maracay (SOLANO, 1969) and Rancho

Grande (RIvERO, 1961).

Other literature records include: (1) the Distrito Federal: Caracas (BOETTGER, 1892;

SoLano, 1969), Mamo, near La Guaira (LUTZ, 1927a-b, 1955), El Limôn, Carayaca and

Cerro Avila (RIVERO, 1961); (2) the Estado Falcôn: Distrito Acosta, Cerro Cosme, Pauji

(SHREVE, 1947; RiveRo, 1961); (3) the Estado Portuguesa: La Apariciôn, between rios Are

Source : MNHN, Paris

112 ALYTES 14 (3)

and Guache (Rivero, 1961, 1964a). New records for Estados Barinas, Carabobo, Falcon,

Portuguesa, and Trujillo, in Venezuela, are listed above.

Populations of the species from the Falcon region of Venezuela, Cordillera de La

Costa and Cordillera de Mérida may currently be isolated from each other by the Yaracuy

Depression and some low xerophytic passes. The areas they currently occupy may be the

product of a formerly wider distribution during previous warmer interglacial periods of the

Quaternary (in which humid forests of all three mountain ranges may have had a more

continuous coverage), and its subsequent humid forest breakdown in drier periods (like

glacial episodes). Many areas in the Lara-Falcon mountain system are good prospective

places where to look for the species. The distribution of B. sternosignatus southward

reaches Barinitas, in the Los Llanos versant of the Cordillera de Mérida; it may be

expected from similar environments down to the Tâchira Depression.

Bufo sternosignatus has been reported from elevations of 150 m (ULABG 3947; see

above) to 1800 m (precise specimen and locality not stated, in FROST, 1985), in humid

premontane forests (“Bosque hümedo premontano” and “Bosque muy hümedo premon-

tano”, in the Holdridge’s Life Zone System for Venezuela of EweL et al., 1976). Most of

the original vegetation in these environments is lost, specially due to its having been turned

into coffee and cacao plantations. The mean annual precipitation of these forests ranges

from 1100 to 4000 mm, with water balance favourable during most part of the year,

although in the Cordillera de La Costa a short dry period occurs (EWEL et al., 1976). In

places with a strong seasonal precipitation (like some forests in the Cordillera de La

Costa), populations of Bufo sternosignatus may behave as explosive breeders. Bufo

sternosignatus, like some other species associated with humid premontane forests (e.g.

Mannophryne spp.; see LA MaARCA, 1992b, 1993, 1994a-b), is most probably restricted to

areas with temperatures between 18 and 24°C. Temperatures away from this range may

constitute a physiological hindrance to the species, because more than 24°C increases the

evapotranspiration rate and less than 18°C increases the probability of frost (EWEL et al.,

1976), thereby resulting in topoclimatic changes that may be adverse to these amphibians.

No ecological data have been gathered on the species, and our only data on prey items

comes from MCNG 2191 and ULABG 2940, which contained ants and beetles (some

Curculionidae).

RESUMEN

En este trabajo proveemos una redescripciôn de Bufo sternosignatus, un sapo pequeño

del noroeste de Venezuela generalmente asociado con el “grupo de B. typhonius”, y datos

detallados sobre morfologia externa, piel, datos meristicos, patrôn de coloracién de

animales vivos y preservados, y biogeografia.

Source : MNHN, Paris

La MarcA & MUARES-URRUTIA 113

ACKNOWLEDGMENTS

The senior author thanks Hobart M. SMITH (University of Colorado), for providing pertinent

literature as well as extending many other courtesies, and Arnold G. KLUGE and Dennis M. HARRIS

for UMMZ computer lists. The junior author thanks Alexis ARENDS, head of the Falcon

Herpetological Survey Project, and Jaime ARANGUREN, Allan L. MARKEZICH and Edgar WEFFER for

their kind field companionship. We acknowledge Amelia Diaz DE PascuAL (CVULA) for loan of

specimens under her care. This paper profited by partial funding through grants CDCHT-ULA

C-564-92A (from the Consejo de Desarrollo Cientifico, Humanistico y Tecnolôgico, Universidad de

los Andes), and S1 9112-030 (from FUNDACITE Falcôn). Drawings were made under a Wild MSAPO

stereomicroscope with camera lucida attachment, acquired through grant S1-2053 from the

Venezuelan Consejo Nacional de Investigaciones Cientificas y Tecnolôgicas (CONICIT). Two

anonymous reviewers provided helpful comments on the manuscript.

LITERATURE CITED

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Naturforschenden Gesellschaft in Frankfurt am Main. Frankfurt am Main, Knauer: i-x + 1-73.

BOULENGER, G. A., 1882. — Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the

British Museum. London, Taylor & Francis: i-xvi + 1-503, pl. I-XXX.

Cat, J. M., 1972. — Bufo of South America. /n: W. F. BLAIR (ed.), Evolution in the genus Bufo, Austin,

Univ. Texas Press: 82-92.

CocHrAN, D. M. & Goïx, C., 1970. — Frogs of Colombia. U.S. natn. Mus. Bull., 288: i-xii + 1-655.

DUuELLMAN, W. E. & SCHÜLTE, R., 1992. — Description of a new species of Bufo from northern Peru

with comments on phenetic groups of South American toads (Anura: Bufonidae). Copeia, 1992

(1): 162-172.

Ewez, J. J., MADRiz, A. & Tosi, J. A., Jr., 1976. — Zonas de Vida de Venezuela. Memoria Explicativa

sobre el Mapa Ecolôgico. 2" ed. Caracas, MAC, FONAIAP: 1-270, 1 map.

FRosT, D. R. (ed.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst.

Coll. : [i-iv] + i-v + 1-732.

HaRDING, K. A., 1983. — Catalogue of New World amphibians. Oxford, Pergamon Press: i-xv +

1-406.

HEYER, W. R., RAND, A. S., GONÇALVES DA CRUZ, C. A., PEIXOTO, O. L. & NELSON, C. E., 1990. —

Frogs of Boracëia. Arg. Zool., 31 (4): 231-410.

HooGMoED, M. S., 1989. — South American bufonids (Amphibia: Anura: Bufonidae), an enigma for

taxonomists. Zn: X. FONTANET & N. HORTA, Treballs d'ictiologia i herpetologia, Tre. Soc. cat.

Ictiol. Herp., 2: 167-180.

ee 1990. — Biosystematics of South American Bufonidae, with special reference to the Bufo

“ryphonius” group. In: G. PErErs & R. HUTTERER (eds.), Vertebrates in the tropies, Bonn,

Museum Alexander Koenig: 113-123.

KELLOGG, R., 1932. — Mexican tailless amphibians in the United States National Museum. Bull. U.

S. nat. Mus., 160: i-iv + 1-224, pl. 1.

La MarCA, E., 1984. — À saxonomic and systematic revision of the frogs of the Colostethus collaris

group (Anura: Leptodactylidae: Dendrobatinae). Master Thesis, Univ. Nebraska: 1-256.

“5 1989. — A new species of collared frog (Anura: Dendrobatidae: Colostethus) from Serrania de

Portuguesa, Andes of Estado Lara, Venezuela. Amphibia-Reptilia, 10: 175-183.

= 19924. — Catälogo taxonômico, biogeogräfico y bibliogräfico de las ranas de Venezuela.

Cuadernos geogräficos, Mérida, Univ. Los Andes, 9: 1-197,

1992. — Geografia de las ranas andinas de Venezuela. Geographica de Mérida, 1 (1): 25-30.

1993. — Phylogenetic relationships and taxonomy of Colostethus mandelorum (Anura: Dendro-

batidae), with notes on coloration, natural history, and description of the tadpole. Bull.

Maryland herp. Soc., 29 (1): 4-19.

Source : MNHN, Paris

114 ALYTES 14 (3)

= 1994a. — Biogeografia de las ranas del género Mannophryne (Anura: Dendrobatidae) en la

América del Sur. Anuario de Investigaciôn 1991, Mérida, IGCRN-ULA: 43-45.

en 1994b. — Taxonomy of the frogs of the genus Mannophryne (Amphibia: Anura: Dendrobatidae).

Publ. Asoc. Amigos Doñana, 4: 1-75.

es 1995. — Crisis de biodiversidad en anfibios de Venezuela: estudio de casos. /n: M. E. ALONSO

(ed), La biodiversidad neotropical y la amenaza de las extinciones, Cuadernos de Quimica

ecolôgica, Mérida, Univ. Los Andes, 4: 47-70.

LuTz, A., 1927a. — Notas sobre batrachios da Venezuela e da Ilha de Trinidad. Mem. Inst. Oswaldo

Cruz, 20 (1): 35-50, pl. 8-15.

ons 1927. — Notes on batrachians from Venezuela and Trinidad. Mem. Inst. Oswaldo Cruz, 20 (1):

51-65, pl. 8-15.

es 1955. — Notas sobre los batracios de Venezuela y de la isla de Trinidad. Mem. Inst. Oswaldo

Cruz, 23: 85-97, pl. 16-24.

LyncH, J. D., 1979. — The amphibians of the lowland tropical forests. p. 189-215. Jn: W. E.

DUELLMAN (ed.), The South American herpetofauna: its origin, evolution, and dispersal, Mus. nat.

Hist. Univ. Kansas Mon., 7: 1-485.

MÜLLER, P., 1973. — The dispersal centres of terrestrial vertebrates in the Neotropical realm.

Biogeographica, The Hague, Junk, 2: 1-244.

PARKER, H. W., 1935. — The frogs, lizards, and snakes of British Guiana. Proc. zool. Soc. London,

1935: 505-530.

PERACCA, M. G., 1914. — Reptiles et Batraciens de Colombia. Mém. Soc. sci. Neuchâtel, 16-111.

PORTER, K. R., 1964. — Distribution and taxonomic status of seven species of Mexican Bufo.

Herpetologica, 19 (4): 229-247.

RIvERO, J. A., 1961. — Salientia of Venezuela. Bull. Mus. comp. Zool., 126: 1-207, 1 pl.

a 1964a. — Salientios (Amphibia) en la colecciôn de la Sociedad de Ciencias Naturales La Salle

de Venezuela. Caribb. J. Sci., 4 (1): 297-305.

= 1964b. — The distribution of Venezuelan frogs. IV. The coastal range. Caribb. J. Sci., 4 (1):

307-319.

--- 1967. — Anfibios coleccionados por la expediciôn franco-venezolana al Alto Orinoco 1951-1952.

Caribb. J. Sci., 7 (3-4): 145-154.

SHREvE, B., 1947. — On Venezuelan reptiles and amphibians collected by Dr. H. G. Kugler. Bull.

Mus. comp. Zool., 99 (5): 519-537.

SmirH, H. M. & TAYLOR, E. H., 1950. — Type localities of Mexican reptiles and amphibians. Univ.

Kansas Sci. Bull., 33 (8): 313-380.

SOoLANO, H., 1969. — Beiträge zur Kenntnis der Amphibienfauna Venezuelas. Vrüff. zool.

Staatssamml. München, 13: 1-26.

WERNER, F., 1916. — Bemerkungen über einige niedere Wirbeltiere der Anden von Kolumbien mit

Beschreibungen neuer Arten. Zool. Anz., 47 (11): 301-304.

Corresponding editor: Jaime PÉFAUR.

Source : MNHN, Paris

Alytes, 1996, 14 (3): 115-121. 115

Acanthocephala parasitic in North American

amphibians: a review with new records

Donald F. MCALPINE

Natural Sciences Department, New Brunswick Museum,

277 Douglas Avenue, Saint John, New Brunswick, E2K 1ES, Canada,

and Biology Department, University of New Brunswick, Fredericton, New Brunswick, E3B 6E1, Canada

Sixty-nine records of Acanthocephala in North American amphibians are

reviewed. In North America, nine adult and two cystacanth Acanthocephala

species in six genera have been recorded from 29 amphibian host species.

Most reports are from aquatic hosts. Acanthocephala in four of these genera

are considered accidental infections. Cystacanths of Centrorhynchus species

and certain adult Fessisentis species are the only regular acanthocephalan

parasites of North American amphibians. New host records for Fessisentis

friedi in Rana pipiens and R. catesbeiana and Neoechinorhynchus rutili in R.

catesbelana collected in New Brunswick, Canada, are additional to the

previous single report of an adult acanthocephalan parasitic in a North

American anuran. In North America, larval Acanthocephala occur in both

salamanders and frogs. Adult Acanthocephala in North American amphibians

most frequently parasitize aquatic salamanders and only parasitize anurans

accidentally. This is in contrast to European amphibians where adult Acan-

thocephalus ranae is common in both salamander and frog helminth commu-

nities and acanthocephalan cystacanths have been infrequently recorded as

amphibian parasites.

Acanthocephalans are infrequent and relatively uncommon parasites of North

American amphibians, although several species, most notably Acanthocephalus ranae

(Schrank, 1788), are common to such hosts in Europe. In this note, I review the scattered

reports of Acanthocephala from amphibians in North America and present new records

of Acanthocephala parasitic in bullfrogs, Rana catesbeiana, and leopard frogs, R. pipiens,

collected in New Brunswick, Canada. Acanthocephalan specimens on which new records

are based have been deposited in the collections of the New Brunswick Museum.

The earliest report of Acanthocephala parasitic in North American Amphibia is that

of Srices & HASssALL (1894), who recorded Echinorhynchus sp. from Notophthalmus

viridescens, the red-spotted newt, collected in Maryland. VAN CLEAVE (1915) later

identified the seven specimens in this collection as Acanthocephalus ranae, a species typical

of European amphibians. MOCALPINE (in press) has re-identified this material as

Acanthocephalus dirus (Van Cleave, 1931). HoLL (1932) commonly encountered an

acanthocephalan in the aquatic adult stage of the red-spotted newt which was subse-

quently described by VAN CLEAVE (1931) as 4. acutulus. MCALPINE (in press) demon-

strated that these worms belong to the genus Fessisentis and should be referred to as

Source : MNHN, Paris

Table I. - Records of Acanthocephala from North American Amphibia. The use of the terms prevalence, intensity (range), and abundance follow MARGOLIS et al.

(1982). Where reported sample size for hosts is shown in parenthesis. Age class of hosts is noted as larvae (*), larvae and adults (*+) or adults (+)

i Intensity*/

Species Age class Host Location Prevalence | lmensie Source

Acanthocephala sp cysacanth Rana syivatica + Ohio 7 7 OLAUG (1954)

Acanthocephala sp cystacanth Desmognathus brimleyorum+ (13) Arkansas 8% 1 WANTER et al. (1986)

Acanthocephala sp ystacanth Desmognathus brimleyorum + (41) Arkansas 28 1 MCALLISTER et a, (1995a)

Acanthocephala sp cystacanth Desmognathus fuscus + (16) North Carolina 31% 137 MANN (1932)

Acanthocephale sp cystacanth Desmognathus fuseus + (16) North Carolina 21% 03° MANN (1932)

Acanthocephala sp cystacanth Desmognaihus fuscus + North Carolina 301% 004 RANKIN (1937)

Acanhocephala sp cystacanh Desmognaihus fuscus + North Carolina 303% 0.06* RANKIN (1937)

Acanthocephala sp cysucanh | Desmognathus quadramaculatus + (46) | North Carolina 0.02% 0.06* RANKIN (1937)

Acanthocephala sp cystacanth Plethodon albagula + (37) Arkansas 30% 1 MCALLISTER (1993)

Acanthocephala sp cystacanth Plethodon glutinosus + (20) North Carolina 50% 020* MANK (1932)

Acanthocephala sp cystacanth Plethodon glutinosus + (39) North Carolina 25% 0.10* MANN (1932)

Acanthocephala sp. adult Ambystoma opacum® (18) North Carolina 312% 0.62* RANKIN (1937)

Acanthocephala sp adult Desmognathus fuscus* North Carolina 49% 0.09 RANKIN (1937)

Acanthocephala sp. adult Desmognalhus fuscus + North Carolina 54% 0.07 RANKIN (1937)

Acanthocephala sp dut Desmognaihus fuseus + (442) Ilinois 11% 001 DvR et al. (1980)

Acanthocephala sp. adut Eurycea muliplicata + Arkansas 71% 2 FOGLE (1960)

Acanthocephala sp. adult Notophthalmus viridescens + North Carolina 31% ou RANKIN (1937)

Acanthocephala sp. adult Plethodon glutinosus + North Carolina 34% 0.06* RANKIN (1937)

Acanhocephala sp. adult Plethodon jordani + (195) North Carolina 12% 1 Dem (1983)

Acanthocephala sp. adult Siren intermedia+ (2) Missouri 50.0% 5 DYER & RANDON (1973)

Acanthocephalus sp. dut Ambystoma talpoideum + (2) Ilinois 50.0% s LANDEVE (1963)

DRE au Anphia ridneyum+ (85) Lousans 12% ie BEN HUNES (939,

Acanthocephalus sp. adult Plethodon glutinosus + (67) Louisiana 15% ‘D NickoL (1967, 1969)

Acanthocephalus dirus aduit Notophthalmus viridescens + Maryland ? ? PARTS

Acanthocephalus dirus adut Necturus maculosus + Wisconsin 429% 6 AMI (1985)

‘Acanthocephalus dirus adult Necturus maculosus + Wisconsin 100.0% 1 AMI (1985)

Fessisentis acurulus adut Notophihalmus viridescens + (123) North Carolina 86,8% 0.64 NÉ

Fessisents fessus aduit Ambystoma talpoïdeum + Ilinois 70% ? BuckNER & NICKOL (1979)

Fessisents fessus adut Siren intemedia+ (G) Ilinois 100.0 5° 610) D ES

Fessisents fessus aduit Siren intermedia+ (68) Ilinois 235% a12 ALTG (1967), NicxOL (1972)

Fessisents fessus adult Siren intermedia + Louisiana ? ? NickoL (1972)

Fessisents fessus adult Ilinois 813% 49) NIcKOL (1972)

Fessisentis fessus adult Ilinois 100% ? DUNAGAN & MILLER (1973)

9IT

(€) +1 SHLATV

Source : MNHN, Paris

Table I. (continued)

à eu Intensity*/

Species Age class Host Location Prevalence MES Source

Fessisentis friedi adult Rana catesbeiana+ (12) New Brunswick 83% dé This report

Fessisentis friedi adult Rana pipiens + (101) New Brunswick 30% 1 This report

Fessisentis friedi adult Rana pipiens (100) New Brunswick 1.0% a This report

Fessisentis friedi adult Necturus maculosus + Ohio ? ? BUCKNER & NICKOL (1979)

Fessisentis necturorum adult Ambysioma opacum* Georgia 86.8% 3.9* (1-22) NICKOL & HEARD (1973)

Fessisentis necturorum adult Eurycea bislineaia / longicauda* Georgia 37.5% 13° (1-2) NICKOL & HEARD (1973)

Fessisentis necturorum adult Necturus beyeri+ (11) Louisana 90.9% 3 NICKOL (1967, 1969)

Fessisentis necturorum adult Notophthalmus viridescens + (6) Georgia 16.7% NickOL & HEARD (1973)

Fessiseniis necturorum adult Pseudotriton montanus* Georgia 96.0% 34° (1-13) NICKOL & HEARD (1973)

Fessisentis vancleavi adult Eurycea longicauda+ (14) Arkansas ? ? SALTARELLI (1977)

Fessisenis vancleavi adut Eurycea mulHplicata+ (19) Oklahoma 428 ? MLEWrrz (1956)

Fessisentis vancleavi adult Eurycea muliplicala + (E) Arkansas 15% ? SALTARELLA (1977)

Fessisenis vancleavi adult Eurycea muliplicata*+ Arkansas ? ? BuckNER & NICKOL (1978)

Fessisentis vancleavi adult Eurycea multiplicata + (50) Arkansas 4% qe MCALLISTER et al. (1995b)

Fessisentis vancleavi adult Eurycea tynerensis+ (13) Oklahoma 13.7% 1.5° (1-3) HUGHES & MOORE (1943)

Pomphorhynchus bulbicolli adult Notophthalmus viridescens + (138) Massachusetts 0.7% 0.08* RANKIN (1945)

Leptorhynchoides thecarus adult Ambystoma tridacrylum + (11) Tennessee 100% (6-30) REIBER (1941)

Leptorhynchoides ihecatus aduit Ambystoma tridacrylum + Tennessee ? ? LINCICOME & VAN CLEAVE (1949)

Leptorhynchoides thecatus adult Necturus maculosus + Indiana F + LINCICOME & VAN CLEAVE (1949).

Centrorhynchus sp. cystacanth Bufo fowleri+ (62) North Carolina 1.6% 0.02° BRANDT (1936)

Centrorhynchus sp. cystacanth Pseudacris crucifer+ (60) North Carolina 1.7% 0.02* BRANDT (1936)

Centrorhynchus sp. cystacanth Pseudacris brimleyi+ (55) North Carolina 73% 0.13* BRANDT (1936)

Centrorhynchus sp. cystacanth Rana catesbeiana+ (33) North Carolina 818% 13.9* BRANDT (1936)

Centrorhynchus sp. Cystacanth Rana catesbeiana+ (38) North Carolina 53% 0.13% BRANDT (1936)

Centrorhynchus sp. cystacanth Rana catesbeiana+ (30) Virginia 333% 52° CAMPBELL (1968)

Centrorhynchus sp. Cystacanth Rana catesbeiana+ (69) Texas 2.9% 1 HoLLis (1972)

Centrorhynchus sp. cystacanth Rana clamitans + (29) Virginia 24.1% 64° CAMPBELL (1968)

Centrorhynchus sp. cystacanth Rana sphenocephala + (60) North Carolina 28.3% 2.6* BRANDT (1936)

Centrorhyÿnchus californicus cystacanth Hyla regilla+ California 46% 1° MILLZNER (1924)

Centrorkynchus conspectus cystacanth Desmognathus fuscus + (54) Louisana 3.17% 2* NICKOL (1969)

Cenirorhynchus conspectus cystacanth Desmognathus monticola+ (125) North Carolina 1.6% 0.02° GOATER et al. (1987)

Centrorhynchus conspectus cystacanth Desmognathus quadramaculatus+ (115) | North Carolina 17% 0.02° GOATER et al. (1987)

Centrorhynchus conspectus cystacanth Plethodon glutinosus + (67) Louisiana 3.0% 2e NICKOL (1969)

Centrorhynchus wardae adult Rana clamitans + (29) Virginia 34% 0.03* CAMPBELL (1968)

Neocchinornehes sp. saut Siren intemeda® Iinois : : Hz & Duvagan (7)

Neoechinorhynchus rutili adult Rana catesbeiana+ (204) New Brunswick 0.5% #* This report

ANIdTYON

LIT

Source : MNHN, Paris

118 ALYTES 14 (3)

Fessisentis acutulus (Van Cleave, 1931), while Acanthocephalus acutulus reported by

NickoL (1969) is assigned to Acanthocephalus sp. Material identified by RANKIN (1937)

and DYER & BRANDON (1973) as 4. acutulus is no longer extant, and the identity of these

worms is therefore unknown.

The 69 natural infections reported in Table I include encysted cystacanths (from the

mesentery, or rarely muscle) and adult Acanthocephala (from the intestine). Experimental

infections of Macracanthorhynchus ingens (Linstow, 1879) in Rana pipiens and of

Fessisentis fessus Van Cleave, 1931 in Ambystoma opacum and À. tigrinum reported by

MooRE (1946) and BUCKNER & NiCKOL (1979) are not included in Table I. Cystacanths

have been identified as Centrorhynchus sp., C. californicus Millzner, 1924 and C. conspectus

Van Cleave & Pratt, 1940. Centrorhynchus californicus was described by MILLZNER (1924)

from the mesentery of Hyla regilla but has not been reported since. Unfortunately, the

type material appears to no longer exist (personal commmunications from J. R.

LiCHTENFELS, U.S. National Parasite Collection, J. P. DONAHUE, Natural History Museum

of Los Angeles, E. KooLs, California Academy of Sciences and D. B. WAKE, Museum of

Vertebrate Zoology, University of California).

The 25 North American amphibian cystacanth records are from both salamanders

and anurans (56 % vs. 44 %). More than 75 % of these reports are from aquatic hosts.

Although hosts such as Bufo fowleri and Pseudacris species are principally terrestrial

outside the breeding season, amphibian cystacanth hosts are most commonly aquatic

species. In Europe, unlike North America, there are few records of cystacanths parasitizing

amphibians (SCHMIDT, 1985). Several extensive surveys of helminths in European

amphibians have not revealed cystacanths parasitizing such hosts (HRISTOVSKI & LEES,

1973; PROKOPIC & KRIVANEC, 1975).

Nine species of adult Acanthocephala in six genera have been recorded as parasites

in the intestines of North American amphibians. Records for Pomphorhynchus bulbocolli

Linkins in Van Cleave, 1919, Leptorhynchoïdes thecatus (Linton, 1891), Centrorhynchus

wardae Holloway, 1958 and Neoechinorhynchus rutili (Müller, 1780) are accidental

infections in amphibians since these Acanthocephala are normally parasitic in fish or birds.

Prevalences of these species in amphibians were mostly under 1 % and reports were often

based on single or few worms. In addition, reports of C. wardae in R. clamitans, L.

thecatus in Amphiuma tridactylum and N. rutili in R. catesbeiana are based on worms

which were not sexually mature. Acanthocephalus dirus infections in Necturus maculosus

appear peripheral to the usual infections in numerous piscine hosts. Although prevalences

of infection were high, host sample sizes for these reports are too small on which to base

any conclusions.

CAMPBELL (1968) provided the only previous North American report of an adult

acanthocephalan in an anuran, Centrorhynchus wardae in Rana clamitans. RICHARDSON

(1993) noted the strong resemblance between C. wardae and C. conspectus. The host

records for F. friedi in R. catesbeiana and R. pipiens presented here are the first reports of

Fessisentis sp. parasitizing Anura. Among amphibians, aquatic salamanders are the most

frequent hosts for Fessisentis species (NickoL, 1969, 1972; NicKOL & HEARD, 1973;

BUCKNER & NICKOL, 1979). In nine of 18 infections of amphibians with Fessisentis species,

where information on prevalences is provided, rates are 70 % or greater, and Fessisentis

Source : MNHN, Paris

MCALPINE 119

vancleavi and F. necturorum are only known as parasites of salamanders (AMIN, 1980).

Nonetheless, species of Fessisentis have not been encountered as widely in North American

amphibians as has been À. ranae in European amphibians.

More than 80 % of North American adult acanthocephalan infections have been

reported in salamanders from aquatic habitats. Similarly, PEARSE (1932) found Acantho-

cephala in Japan restricted to salamanders from aquatic environments, suggesting this was

because of a dependence on aquatic intermediate hosts. NickoL & HEARD (1973) reported

that F. necturorum parasitized only aquatic stages of salamander hosts. Adult Acantho-

cephala appear rarely to infect North American anurans or any terrestrial or semi-

terrestrial amphibian. The accidental infection in R. clamitans reported by CAMPBELL

(1968), and the Acanthocephala records for R. catesbeiana and R. pipiens reported here,

are the only cases of adult Acanthocephala parasitic in North American frogs. In contrast,

A. ranae has been regularly reported from both European frogs and salamanders (LEES,

1962; PRoKkoPIC & KRIVANEC, 1975; KuC & SULGOSTOWSKA 1988a), sometimes at

prevalences of over 80 % (Kuc & SuLGosTowskA 1988b). WALTON (1942) noted that N.

rutili has been reported parasitic in R. esculenta from Europe.

Although AO (1990) noted that few parasite surveys of amphibians have been

comprehensive, it appears that helminth communities of both European salamanders and

frogs regularly include adult Acanthocephala, whereas North American amphibian

helminth communities do so infrequently.

ACKNOWLEDGEMENTS

Acanthocephalans reported here were collected during a survey of the helminths of bullfrogs,

green frogs and leopard frogs in New Brunswick. I am grateful to T. J. FLETCHER, C. WAGG, K.

RouRkE and T. GoMar for assistance in collecting frogs.

LITERATURE CITED

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120 ALYTES 14 (3)

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Source : MNHN, Paris

MCALPINE 121

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Source : MNHN, Paris

Alytes, 1996, 14 (3): 122-126.

Helminths of the oak toad

(Bufo quercicus, Bufonidae)

from Florida (U.S.A.)

Stephen R. GOLDBERG * & Charles R. BURSEY **

* Department of Biology, Whittier College, Whittier, California 90608, U.S.A.

** Department of Biology, Pennsylvania State University, Shenango Valley Campus, 147 Shenango Avenue,

Sharon, Pennsylvania 16146, U.S.A.

The gastrointestinal tracts, lungs and urinary bladders of 35 Bufo querci-

cus from Florida (U.S.A) were examined for helminths. Four genera of

nematodes (Ascarops, Cosmocercoides, Filariidae gen. indet. and Physalo-

ptera) and one genus of acanthocephalan (Polymorphus) were found. The

highest prevalence was 11% for Cosmocercoides, and the greatest mean

intensity was 4.5 for Ascarops sp. Bufo quercicus is a new host record for each

of these genera of helminths.

The oak toad, Bufo quercicus Holbrook, 1840, occurs in pine woods of the southern

coastal plain from southeast Virginia throughout Florida to eastern Louisiana (CONANT &

CoLLins, 1991). It is the smallest bufonid in the United States, reaching a maximum size

of only 38 mm (AsHTON & ASHTON, 1988). To our knowledge, there are only two reports

of helminths in oak toads (WALTON, 1938; HAMILTON, 1955). With worldwide attention

focussed on declining amphibians (HEYER et al., 1994), knowledge of the occurrences of

helminths and their possible negative impact on anurans has become a topic of interest.

The purpose of this paper is to report the occurrences of helminths in a Florida population

of Bufo quercicus as part of an ongoing study of the biogeography of North American

bufonid helminths.

Thirty five Bufo quercicus from Florida, U.S.A. (30 males, 5 females, mean snout-vent

length SVL = 27.9 mm + 2.8 SD, range 24-35 mm) were borrowed from the herpetology

collection of the University of Florida, Gainesville: UF 9602.1-19, collected April, 1957,

Palm Beach County; UF 66801-66810, collected July 1970, Leon County; UF 100225-

100230, collected May-June 1977 or 1978, Marion County.

The toads were dissected in situ, i.e. without removing organs from the body. The

body cavity was opened by a longitudinal incision from throat to vent. The lungs,

esophagus, stomach, small intestine, large intestine and bladder were examined under a

dissecting microscope. AI helminths were removed and identified using a standard glycerol

wet mount procedure. Terminology is in accordance with MARGOLIS et al. (1982).

Source : MNHN, Paris

GoLDBERG & BURSEY 123

Ten (29 %) of the 35 Bufo quercicus harbored helminths. Nematodes included five

adult females of Cosmocercoides sp., three third-stage larvae of Physaloptera sp., nine

encapsulated larvae of Ascarops sp., two third-stage larvae of Filariidae gen. indet. and

two acanthocephalan cystacanths, Polymorphus sp. Helminths were found in the stomach,

large intestine and coelom; none were found in the lungs, esophagus, small intestine or

bladder. Voucher specimens were deposited in the United States National Parasite

Collection, Beltsville, Maryland 20705, U.S.A.: Ascarops sp. (85877-85878), Cosmocercoi-

des sp. (85879-85881, 85886); Physaloptera sp. (85876, 85882, 85885); Filariidae (85884),

Polymorphus sp. (85883, 85887).

One female and two male toads from Leon County and one female from Marion

County harbored five adult females of the genus Cosmocercoides (prevalence: 11 %). This

represents the highest prevalence found for a helminth in our study. These nematodes were

found in the large intestines. Since male nematodes were absent from the collection, it was

not possible to determine the species. Cosmocercoides dukae has previously been reported

from Bufo quercicus (WALTON, 1938); however, the collection locality was not stated.

Cosmocercoides variabilis (previously known as Oxysomatium variabilis) was found in B.

quercicus from Florida and/or Georgia (HAMILTON, 1955). Only two species of Cosmo-

cercoides have been reported from North America, namely C. dukae and C. variabilis

(BAKER, 1987). These two species have caused some taxonomic problems in that C.

variabilis was at one time considered to be a synonym of C. dukae. However,

VANDERBURGH & ANDERSON (1987a-b) showed that such synonymy was incorrect and

reported C. variabilis to be a normal parasite of the Bufonidae, while C. dukae is normally

a parasite of molluscs but may occur accidentally in frogs and salamanders which feed on

molluscs (VANDERBURGH & ANDERSON, 1987c). Thus, the specimens recorded by WALTON

(1938) should be referred to C. variabilis.

One female and one male toads from Marion County and one female from Palm

Beach County harbored three third-stage larvae of Physaloptera sp. (prevalence: 9 %).

Two larvae were found in stomachs, one was found in the large intestine. Larval

physalopterans have been found in several species of Bufo, namely, B. alvarius, B.

americanus, B. cognatus, B. debilis, B. microscaphus, B. retiformis, B. speciosus (as B.

compactilis) and B. woodhousii (KUNTZ, 1940; PARRY & GRUNDMANN, 1965; ASHTON &

RABALAIS, 1978; GOLDBERG & BURSEY, 1991; GoLDBERG et al., 1995, 1996), but,

apparently, no cases of parasitism of toads by adult physalopterans have been reported.

The presence of larvae and lack of adult physalopterans has been reported in many other

species of amphibians and reptiles (see GOLDBERG et al., 1993). Species of Physaloptera

require an insect intermediate host (ANDERSON, 1992); thus, the presence of their larvae in

insectivorous species is not unexpected. Because physalopterines are usually found

attached to the gastric mucosa (ANDERSON, 1992), our finding one larva in the large

intestine suggests that toads are not appropriate hosts for this species and development to

maturity does not occur in them.

Two male toads from Palm Beach County (prevalence: 6%) harbored nine

encapsulated larvae of Ascarops sp. for a mean intensity of 4.5, highest found in our study.

The cysts occurred in the stomach wall, two in one toad and seven in the other.

Encapsulated larvae of Ascarops sp. have been found in lizards (GOLDBERG & BURSEY,

Source : MNHN, Paris

124 ALYTES 14 (3)

1988, 1989; MCALLISTER et al., 1993), birds (KRAHWINKEL & MCCUE, 1967) and mammals

(ALICATA & MCINTOSH, 1933; CHANDLER, 1946), but this is apparently the first record of

this genus in toads. Two hosts are required for development: adults are found in the

stomach of swine, rats and mice, in the esophagus of ruminants and in the crop of chickens

(OLSEN, 1974); larvae develop in Coleoptera and Odonata (ALICATA, 1935). Ascarops sp.

larvae can be expected in animals that habitually feed upon insects.

One female toad from Marion County (prevalence: 3 %) harbored two filarial larvae

in the coelom. Since the reproductive system had not begun to develop, no attempt was

made to identify the specimens. Apparently no adult filariids have been reported from

North American toads; but, species of Foleyellides do parasitize Rana utricularia

sphenocephala in Florida (BAKER, 1987). Whether filariids actually parasitize North

American toads or those found in this study represent an accidental infection is not

presently known. Further work will be required to answer this question.

One female and one male toads from Marion County (prevalence: 6 %) harbored two

acanthocephalan cystacanths. They were in the coelom but attached to the stomach wall.

These cystacanths were orange in color, about 0.5 mm in length, and appeared to lack

genital spines. For these reasons the cystacanths were identified as Polymorphus sp.

Amphipods serve as intermediate hosts and ducks serve as definitive hosts for species of

Polymorphus (PoDESTA & HOLMES, 1970). To our knowledge, there are no other reports of

cystacanths in North American toads.

WALTON (1938) found one additional nematode from Bufo quercicus, namely

Oswaldocruzia pipiens. Later, HAMILTON (1955) reported Oswaldocruzia subauricularis from

Bufo quercicus. BAKER (1977) reviewed the North American species of Oswaldocruzia and

concluded that a single species, O. pipiens, was present in North American toads and frogs.

BAKER (1987) considered O. subauricularis to be a South American species; thus, the

HAMILTON (1955) record should be referred to as O. pipiens.

Currently, Bufo quercicus may be listed as definitive host of two species of nematodes,

Cosmocercoides variabilis and Oswaldocruzia pipiens, but may serve as a paratenic or

accidental host for several other helminths. Prevalence is apparently low: 29 % in this

study, 7 % in the study by HAMILTON (1955). Because only small numbers of specimens are

available from museums for dissections, our sample was restricted to 35 Bufo quercicus.

Examination of larger samples from throughout the range of Bufo quercicus may yield

helminth species not previously found.

In addition to Bufo quercicus, the Floridian bufonid fauna consists of two native

(Bufo terrestris and Bufo woodhousii fowleri) and one introduced species (Bufo marinus)

(ASHTON & ASHTON, 1988). Apparently, there are no reports of helminths in these species

in the state. Subsequent helminthological examinations will be required before the

helminth fauna of these Floridian toads can be compared.

ACKNOWLEDGMENTS

We thank D. L. AuTH, Division of Herpetology, Florida Museum of Natural History, University

of Florida, for permission to examine Bufo quercicus.

Source : MNHN, Paris

GOLDBERG & BURSEY 125

LITERATURE CITED

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AsHTon, R. E., Jr. & ASHTON, P. S., 1988. — Handbook of reptiles and amphibians of Florida, Part

3, The amphibians, Miami, Windward Publishing, In -191.

BAKER, M. R., 1977. — Redescription of Oswaldocruzia pipiens Walton, 1929 (Nematoda:

Trichostrongylidae) from amphibians of eastern North America. Can. J. Zool., 55: 104-109.

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CHANDLER, A. C., 1946. — Helminths of armadillos, Dasypus novemcinctus, in eastern Texas. J.

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ConanT, R. & COLLINS, J. T., 1991. — A field guide to reptiles and amphibians. Eastern and central

North America. Boston, Houghton Mifflin Company: 1-450.

GoLDBerG, S. R. & BURSEY, C. R., 1988. — Larval nematodes (Ascarops sp., Spirurida, Spirocercidae)

in liver granulomata of the western fence lizard, Sceloporus occidentalis (Iguanidae). J. Wild.

Dis., 24: 568-571.

1989. — Larval nematodes (Ascarops sp.) in stomach granulomas of the sagebrush lizard,

Sceloporus graciosus. J. Wild. Dis., 25: 630-633.

Ep 1991. — Helminths of three toads, Bufo alvarius, Bufo cognatus (Bufonidae), and Scaphiopus

couchii (Pelobatidae), from southern Arizona. J. helm. Soc. Washington, 58: 142-146.

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sympatric toad species, Bufo cognatus, Bufo debilis (Bufonidae), and Spea multiplicata

(Pelobatidae) from New Mexico. J. helm. Soc. Washington, 62: 57-61.

GOLDBERG, S. R., BURSEY, C. R., SULLIVAN, B. K. & TRUONG, Q. A., 1996. — Helminths of the

Sonoran green toad, Bufo retiformis (Bufonidae), from southern Arizona. J. helm. Soc.

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GoLDBERG, S. R., BURSEY, C. R. & TAwiL, R., 1993. — Gastrointestinal helminths of the western

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92: 43-51.

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HEYER, W. R., DONNELLY, M. A., MCDIARMID, R. W., HAYEK, L. C. & FOSTER, M. S. (eds.), 1994.

— Measuring and monitoring biological diversity. Standard methods for amphibians. Washington,

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KRAHWINKEL, D. J., Jr. & MCCUE, J. F., 1967. — Wild birds as transport hosts of Spirocerca lupi in

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KunTz, R. E., 1940. — A study of anuran parasites of Comanche County, Oklahoma. M.S. Thesis,

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MarGoLis, L., EscH, G. W., HOLMEs, J. C., KuRIs, A. M. & SCHAD, G. A., 1982. — The use of

ecological terms in parasitology (report of an ad hoc committee of the American Society of

Parasitologists). J. Paras., 68: 131-133.

MCALLISTER, C. T., GOLDBERG, S. R., BURSEY, C. R., FREED, P. S. & HOLSHUH, H. J., 1993. — Larval

Ascarops sp. (Nematoda: Spirurida) in introduced Mediterranean geckos, Hemidactylus turcicus

(Sauria: Gekkonidae), from Texas. J. helm. Soc. Washington, 60: 280-282.

Source : MNHN, Paris

126 ALYTES 14 (3)

OLSEN, O. W., 1974. — Animal parasites. Their life cycles and ecology. Baltimore, University Park

Press: 1-562.

PARRY, J. E. & GRUNDMANN, A. W., 1965. — Species composition and distribution of the parasites

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occurring as juveniles in Hyalella azteca (Amphipoda) at Cooking Lake, Alberta. J. Paras., 56:

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VANDERBURGH, D. J. & ANDERSON, R. C., 1987a. — The relationship between nematodes of the genus

Cosmocercoides Wilkie, 1930 (Nematoda: Cosmocercoidea) in toads (Bufo americanus) and

slugs (Deroceras laeve). Can. J. Zool., 65: 1650-1661.

1987b. — Preliminary observations on seasonal changes in prevalence and intensity of

Cosmocercoides variabilis (Nematoda: Cosmocercoidea) in Bufo americanus (Amphibia). Can.

J. Zool., 65: 1666-1667.

Es 1987c. — Seasonal changes in prevalence and intensity of Cosmocercoides dukae (Nematoda:

Cosmocercoidea) in Deroceras laeve (Mollusca). Can. J. Zool., 65: 1662-1665.

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Corresponding editors: Janalee P. CALDWELL & Alain DuBois.

Source : MNHN, Paris

Alytes, 1996, 14 (3): 127-128. 127

Erratum

In page 131 of the article quoted below, an unfortunate mistake has led to publication

of a version of figure 7 that does not fit with its legend. This earlier version of the figure

showed diagrammatic representations of larval tooth row formulae of only 9 species of the

genera Osteocephalus and Phrynohyas, and did not include detailed explanations for the

signs and symbols used in the figure. On the other hand, the accompanying legend was

that of the final version of this figure, showing 10 species and including detailed

explanations of the notation system used in this figure. This is all the more annoying as

this notation system was new, and may become adopted by other colleagues for the

schematic representation of tadpoles’ mouthparts. Next page, we present the figure in its

final version, as it should have appeared in the original paper. In order to allow

subsequent readers to trace this erratum, we suggest to quote this paper in the future as

follows:

ScHiEsai, L. C., GRILLITSCH, B. & VocL, C., 1996. — Comparative morphology of phytotelmonous

and pond-dwelling larvae of four neotropical treefrog species (Anura, Hylidae, Osteocephalus

oophagus, Osteocephalus taurinus, Phrynohyas resinifictrix, Phrynohyas venulosa). Alytes, 13 (4):

109-139; 14 (3): 127-128.

Alain DuBois

Chief Editor of Alytes

Source : MNHN, Paris

128 ALYTES 14 (3)

O O O

O. oophagus

©. verruciger

P. coriacea P. mesophaea P. resinifictrix

o Beak

Toolh rows present

En, in ectotrophic larval stages,

RS at least in stages 38 + 2,

or absent in stages 38 + 2,

ë Le without further information,

‘ . frequently broken, poorly formed

P. venulosa

Fig. 7. — Variation of larval tooth row formulae among Osteocephalus and Phrynohyas species.

Schematic drawings; median interruptions and relative lengths of lower tooth rows not

considered.

References: Osteocephalus buckleyi (HERO, 1990); Osteocephalus elkejungingerae (HENLE,

1981); Osteocephalus langsdorffii (DUELLMAN, 1974); Osteocephalus oophagus (present study and

as in Table 1); Osteocephalus taurinus (present study and as in Table 1); Osteocephalus verruciger

(TRUEB & DUELLMAN, 1970); Phrynohyas coriacea (SCHIESARI & MOREIRA, in press); Phrynohyas

mesophaea (LUTZ, 1973; SCHIESARI, personal observation); Phrynohyas resinifictrix (present study

and as in Table 1); Phrynohyas venulosa (present study and as in Table 1).

Source : MNHN, Paris

AUVTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1996

Chief Editor: Alain Dusois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

naturelle, 25 rue Cuvier, 75005 Paris, France).

Deputy Editor: Janalee P. CALDWELL (Oklahoma Museum of Natural History, University of Oklahoma,

Norman, Oklahoma 73019, U.S.A.).

Editorial Board: Jean-Louis ALBARET (Paris, France), Ronald G. ALTIG (Mississippi State University,

U.S.A.); Emilio BALLETTO (Torino, Italy); Alain COLLENOT (Paris, France); Günter GOLLMANN (Wien,

Austria); Tim HaLuiDay (Milton Keynes, United Kingdom), W. Ronald Hever (Washington,

U.S.A.}; Walter HôDL (Wien, Austria); Pierre JoLY (Lyon, France); Masafumi Marsui (Kyoto,

Japan), Jaime E. PÉFauR (Mérida, Venezuela); J. Dale Rogers (Perth, Australia); Ulrich SINSCH

(Koblenz, Germany); Marvalee H. WAKE (Berkeley, U.S.A.).

Technical Editorial Team (Paris, France): Alain DuBois (texts); Roger BOUR (tables); Annemarie

OuLER (figures).

Index Editors: Annemarie OHLER (Paris, France); Stephen J. RICHARDS (Townsville, Australia).

GUIDE FOR AUTHORS

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with

amphibians. Beside articles and notes reporting results of original research, consideration is given for

publication to synthetic review articles, book reviews, comments and replies, and to papers based upon

original high quality illustrations (such as color or black and white photographs), showing beautiful or rare

species, interesting behaviors, etc.

The title should be followed by the name(s) and address(es) of the author(s). The text should be

typewritten or printed double-spaced on one side of the paper. The manuscrit should be organized as

follows: English abstract, introduction, material and methods, results, discussion, conclusion, French or

Spanish abstract, acknowledgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or Table IV. Figures should not

exceed 16 x 24 cm. The size of the lettering should ensure its legibility after reduction. The legends of figures

and tables should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRAF & POLLS PELAZ, 1989;

INGER et al., 1974). References in the literature cited section should be presented as follows:

BouRRET, R., 1942. — Les Batraciens de l'Indochine. Hanoï, Institut Océanographique de l'Indochine: i-x

+ 1-547, pl. I-IV.

GRAF, J.-D. & POLLS PELAz, M., 1989. — Evolutionary genetics of the Rana esculenta complex. In: R. M.

DawWLey & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The New York

State Museum: 289-302.

INGER, R. F., Voris, H. K. & Voris, H. H., 1974. — Genetic variation and population ecology of some

Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet, 12: 121-145.

Manuscripts should be submitted in triplieate either to Alain Durois (address above) if dealing with

amphibian morphology, systematics, biogeography, evolution, genetics or developmental biology, or to

Janalee P. CALDWELL (address above) if dealing with amphibian population genetics, ecology, ethology or

life history. Acceptance for publication will be decided by the editors following review by at least two

referees.

If possible, after acceptance, a copy of the final manuscrit on a floppy disk (3 or 5 %) should

be sent to the Chief Editor. We welcome the following formats of text processing: (1) preferably, MS Word

{1.1 to 6.0, DOS or Windows), WordPerfect (4.1 to 5.1, DOS or Windows) or WordStar (3.3 to 7.0); (2) less

preferably, formated DOS (ASCII) or DOS-formated MS Word for the Macintosh (on a 3 ‘4 high density

1.44 Mo floppy disk only).

No page charges are requested from the author(s), but the publication of color photographs is

charged. For each published paper, 25 free reprints are offered by Alytes to the author(s). Additional reprints

may be purchased.

Published with the support of AALRAM

(Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum national d'Histoire naturelle, Paris, France).

Directeur de la Publication: Alain DuBois.

Numéro de Commission Paritaire: 64851.

Alytes, 1996, 14 (3): 101-128.

Contents

Enrique LA MaRCA & Abraham MHARES-URRUTIA

Taxonomy and geographic distribution of a northwestern

Venezuelan toad (Anura, Bufonidae, Bufo sternosignatus) .......... 101-114

Donald F. MCALPINE

Acanthocephala parasitic in North American amphibians:

A TÉVIEWAWILRENEW I TECOIAS nee eee eee eee 115-121

Stephen R. GoLDBERG & Charles R. BURSEY

Helminths of the oak toad (Bufo quercicus, Bufonidae)

OM EION ANUS A) EEE REP eee cnrs 122-126

Erratum

Luis C. ScHisARI, Britta GriLLirscH & Claus VOGL

Comparative morphology of phytotelmonous and pond-dwelling

larvae of four neotropical treefrog species (Anura, Hylidae,

Osteocephalus oophagus, Osteocephalus taurinus, Phrynohyas

HeSHHICINEX, BRTVHORYAS SV ENUIDSA). Re cerner 127-128

Alytes is printed on acid-free paper.

Alytes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 4° trimestre 1996.

Source : MNHN, Paris: