Rediscovery, resurrection and redescription of Rhinella leptoscelis (Boulenger, 1912) (Anura: Bufonidae)

Abstract

We resurrect Rhinella leptoscelis (Anura: Bufonidae) from the synonymy of R. veraguensis and redescribe it on the basis of the holotype, and recently collected specimens from Peru and Bolivia. Rhinella leptoscelis is well distinguished from all other species of the R. veraguensis group by its flat head with well developed orbitotympanic and postorbital crests, distinct tympanum, large parotoid glands, spiny tubercles on dorsal surfaces, long and slender extremities, dorsolateral row of conical, enlarged and elevated tubercles, webbing absent on fingers, basal and serrated webbing between toes, and first finger longer than second. This species is known from the humid forests of the Amazonian versant of the Andes from central Bolivia to southern Peru.

Full text

56 Accepted by M. Vences: 29 Apr. 2009; published: 22 May 2009
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Copyright © 2009 · Magnolia Press
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Rediscovery, resurrection and redescription of Rhinella leptoscelis (Boulenger,
1912) (Anura: Bufonidae)
JOSÉ M. PADIAL1,5, JUAN C. CHAPARRO2, JÖRN KÖHLER3 & IGNACIO DE LA RIVA4
1Department of Evolution, genomics and Systematics, Evolutionary Biology Centre (EBC), Uppsala University, Norbyvägen 18D,
75236 Uppsala, Sweden. E-mail: jose.m.padial@ebc.uu.se
2Museo de Historia Natural, Universidad Nacional de San Antonio Abad del Cusco, Cusco, Peru. Plaza de Armas s/n (Paraninfo
Universitario), Cusco, Peru. E-mail: jchaparroauza@yahoo.com
3Hessisches Landesmuseum Darmstadt, Department of Natural History – Zoology, Friedensplatz 1, 64283 Darmstadt, Germany.
E-mail: joern.koehler@hlmd.de
4Department of Biodiversity and Evolutionary Biology, Museo Nacional de Ciencias Naturales, CSIC; C/José Gutiérrez Abascal, 2,
28006, Madrid, Spain. E-mail: iriva@mncn.csic.es
5Corresponding author
Abstract
We resurrect Rhinella leptoscelis (Anura: Bufonidae) from the synonymy of R. veraguensis and redescribe it on the basis
of the holotype, and recently collected specimens from Peru and Bolivia. Rhinella leptoscelis is well distinguished from
all other species of the R. veraguensis group by its flat head with well developed orbitotympanic and postorbital crests,
distinct tympanum, large parotoid glands, spiny tubercles on dorsal surfaces, long and slender extremities, dorsolateral
row of conical, enlarged and elevated tubercles, webbing absent on fingers, basal and serrated webbing between toes, and
first finger longer than second. This species is known from the humid forests of the Amazonian versant of the Andes
from central Bolivia to southern Peru.
Key words: Amazon, Andes, Peru, Bolivia, Rhinella veraguensis group, Rhinella margaritifera group, taxonomy
Introduction
The taxonomic status of the toad Rhinella leptoscelis (Boulenger, 1912) has remained uncertain. This species,
described on the basis of a single specimen from the Andean slopes of southern Peru, has been considered
either as a junior synonym of R. veraguensis (Schmidt, 1857) (Vellard 1959; Hoogmoed 1990) or as a valid
species (Gallardo 1961; Cei 1968, 1972; Gorham 1974; Hoogmoed 1985; Duellman and Schulte 1992).
Vellard (1959) considered R. ockendeni (Boulenger, 1902) and R. leptoscelis very similar, but recommended
to gather more information to solve the taxonomic status of both species. Savage (1969) synonymized R.
ockendeni under R. veraguensis. Hoogmoed (1990) placed R. leptoscelis in the synonym of R. veraguensis,
and attributed some differences between the single known specimen of R. leptoscelis, a female, and the
lectotypes of R. veraguensis, all adult males, to sexual dimorphism. He furthermore removed R. veraguensis
and all its junior synonyms from the R. typhonius species group and placed it in the R. veraguensis group.
Duellman and Schulte (1992) did not follow this position and treated R. leptoscelis as a valid species without
further discussion. But subsequent taxonomic studies of toads assigned to this group did not considered R.
leptoscelis as a valid species (Harvey and Smith 1993, 1994; Lehr et al. 2001, 2005; Padial et al. 2006;
Pramuk 2006; Chaparro et al. 2007; see account by Frost 2009).
Our comparisons of Rhinella leptoscelis with all type specimens of the R. veraguensis and R.
margaritifera groups inhabiting the Andean slopes of Southern Peru and Bolivia (see De la Riva et al. 2000;

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REDISCOVERY OF RHINELLA LEPTOSCELIS
Köhler 2000; Padial et al. 2006; Chaparro et al. 2007), together with the finding of new specimens at the type
locality (Santo Domingo de Carabaya, Puno, Peru) of R. leptoscelis, indicate that this is a valid species, which
we redescribe and resurrect herein.
Material and methods
For morphological and color characteristics used in the diagnosis and description we follow Duellman and
Schulte (1992). Specimens examined are listed in the Appendix. Measurements were taken with a digital
caliper to the nearest 0.01 mm, but following Hayek et al. (2001), for avoiding pseudo precision, we rounded
all measurements to only one decimal. Abbreviations of measurements are as follows: snout–vent length,
SVL; head length (from posterior margin of the lower jaw to tip of snout), HL; head width, HW; upper eyelid
width, EW; eye diameter (measured horizontally), ED; eye to nostril distance, EN; distance between nostrils,
IND; tibia length, TL; foot length (from proximal border of inner metatarsal tubercle to tip of fourth toe), FL.
Museum acronyms refer to: Natural History Museum, London, U.K. (BM); Centro de Biodiversidad y
Genética, Cochabamba, Bolivia (CBG); Colección Boliviana de Fauna, La Paz, Bolivia (CBF); Museo de
Historia Natural Noel Kempff Mercado, Santa Cruz de la Sierra, Bolivia (MNK-A [Amphibian Collection]);
Estación Biológica de Doñana, Sevilla, Spain (EBD); Museo Nacional de Ciencias Naturales, Madrid, Spain
(MNCN); United States National Museum of Natural History, Smithsonian Institution, Washington, USA
(USNM); and Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK).
We follow Chaparro et al. (2007) in that all South American bufonids assigned to Chaunus by Frost et al.
(2006) should now be included in Rhinell a, except Rhaebo for the former Bufo guttatus group, and
Nannophryne for the former Bufo variegatus group. There seems to be no morphological synapomorphy for
the R. veraguensis group (Pramuk 2006). Phylogenetic analyses (Pramuk 2006; Chaparro et al. 2007) place
some members of the R. veraguensis group as the sister clade of the R. margaritifera group while other
members are basal to it, also suggesting thus that the R. veraguensis group, as defined by Duellman and
Schulte (1992), is paraphyletic. Thus, apart of the general morphological appearance, there is no objective
criterion to definitively assign species to this group. Rhinella leptoscelis is most similar in external
morphology to members of the R. veraguensis group (see below) and to a lesser extent to members of the R.
margaritifera group. We therefore compare R. leptoscelis with members of the R. veraguensis group (listed by
Padial et al. 2006) and with two members of the R. margaritifera group that potentially inhabit the same area
(De la Riva et al. 2000). All those species share the following external characters: medium size; long and
slender extremities; head flattened, wider than long; supraorbital and/or parietal crests prominent; skin on
dorsum and extremities bearing tubercles; lateral row of enlarged tubercles; parotoid glands moderately
developed.
Results
Rhinella leptoscelis (Boulenger, 1912)
Bufo leptoscelis Boulenger, 1912, Ann. Mag. Nat. Hist., Ser. 8, 10: 186. Holotype (Fig. 1A–B): BM 1907.5.7.32. Type
locality: "Santo Domingo, Carabaya, S.E. Peru, 6500 feet".
Diagnosis. A medium-sized species of Rhinella (Figs. 1A–B, 2A–G) distinguished by the following unique
combination of characteristics: (1) canthus rostralis concave in dorsal view, sharp in profile, orbitotympanic
and postorbital crests prominent, not protruding; (2) tympanum distinct, oval; (3) parotoid glands large,
oblong to subtriangular, protuberant, separated from eyelid and tympanum by the supratympanic crest; (4)
body surfaces covered by spiny tubercles with keratinized tips; (5) extremities long and slender; (6) a

PADIAL ET AL.58 · Zootaxa 2115 © 2009 Magnolia Press
dorsolateral row of conical, enlarged, and elevated tubercles; (7) tarsal fold absent; (8) webbing absent on
fingers, basal and serrated between toes; (9) first finger longer than second; (10) iris green in life.
From other species of the Rhinella veraguensis species group, R. leptoscelis can be distinguished as
follows. From R. amboroensis, R. arborescandens, R. justinianoi, R. quechua and R. veraguensis (Fig. 1C–D)
by having a conspicuous tympanic membrane and toe webbing basal and serrated. From R. arborescandens,
R. chavin, R. manu, R. multiverrucosus, R. nesiotes, R. rumbolli, and R. tacana by lacking fleshy interdigital
membranes, having fingers slender with first finger longer than second, and subtriangular parotoids. From R.
chavin and R. multiverrucosus it can be distinguished by lacking glands on the forearms and hind legs.
Rhinella leptoscelis shares with R. fissipes the slender fingers and serrated basal membrane on toes. However,
R. leptoscelis can be distinguished from R. fissipes by having cranial crests, conspicuous tympanic membrane,
and larger tubercles on dorsal skin. With R. inca, R. leptoscelis shares the presence of cranial crests, first
finger longer than second, and the observable tympanic membrane, but R. leptoscelis has more spiny dorsal
skin and serrated fringes, and prominent cranial crests anterior and posterior to eye (absent in R. inca). From
Rhinella margaritifera and R. stanlaii, two members of the R. margaritifera group inhabiting the Andean
slopes from Southern Peru to Central Bolivia, R. leptoscelis can be distinguished by lacking a protruding
cranial crest above the tympanic area.
Redescription of the holotype. An adult female (Fig. 1A–B) with long and slender extremities; large and
flat head, wider than long, its width 36% of SVL, its length 29% of SVL; snout subacuminate in dorsal view,
round in profile; cranial crests present, occipital and orbitotympanic crests well developed but not protruding;
skin of head not co-ossified with underlying cranial bones; internarial area concave; nostrils not protuberant,
very small, oriented laterally; canthus rostralis narrow and elevated in profile, concave in dorsal view; lips
flat; eye-nostril distance smaller than eye length; tympanic membrane visible, conspicuous, oval, surrounded
by tubercles, its diameter approximately half of eye length; tympanic annulus thin, overlapped with
surrounding granules. Forelimbs long and slender; hand broad, with long and slender fingers; relative length
of fingers II<III<I<IV; webbing absent, lateral fringes present, serrated; tips of digits round, not expanded;
ulnar region covered by keratinized spines; palmar tubercle subtriangular, protruding; prepollical tubercle
elongated, protruding, smaller than palmar tubercle; subarticular tubercles round, conspicuous;
supernumerary tubercles conical, diffuse, smaller than subarticular tubercles. Hind limbs and feet long; tibia
length 45% of SVL; foot length 44% of SVL; no tarsal fold, tarsus covered by spines; outer metatarsal
tubercle round, prominent, 1/2 the size of inner; inner metatarsal tubercle small, prominent, elongate; relative
length of toes I<II V<III<IV; webbing basal between toes I–II, II–III, and IV–V, slightly more developed
between III–IV, reaching the basis of second subarticular tubercle of toe III; tip of toes rounded; subarticular
tubercles prominent, round to ovoid, larger than supernumerary tubercles; supernumerary tubercles abundant,
small, conical or rounded. Choanae, vomerine teeth, and tongue conditions were not examined in the holotype
to avoid damaging of the specimen (but see below).
Skin of head, body, and extremities bearing abundant conical tubercles with keratinized tips; enlarged
tubercles forming an irregular dorsolateral row on each side of dorsum; canthus rostralis, interorbital region,
and snout with scarce or no tubercles; parotoid glands large (60% HL), subtriangular, more displaced to side
of the head than to dorsum, almost in contact with tympanic membrane, and in contact with small, short
orbitotympanic and postorbital crest; enlarged glands absent on limbs or forearms; skin on throat and other
ventral surfaces granular to spiny. In preservative, dorsum and ventral surfaces mostly dark reddish-orange,
head, belly and fingers paler.
Measurements (in mm) are only available for adult females. Measurements of the holotype (BM
1947.2.21.95) are followed in parentheses by those of additional specimens (MHNC 5976, 5989, and 5975,
respectively): SVL, 53.5 (85.5, 56.8, 59.8); HL, 15.4 (23.3, 17.4, 19.1); HW, 19.2 (29.3, 20.8, 19.9); ED, 6.5
(6.9, 5.9, 6.9); END, not measured (5.8, 4.2, 5.6); EW, 3.8 (6.0, 4.8, 4.9); IND, not measured (5.2, 3.7, 3.8);
IOD, not measured (10.9, 8.6, 7.0); TL, 24.0 (37.1, 30.4, 28.2); FL, 23.5 (34.5, 27.1, 27.2). These
measurements indicate a large variation in adult female size, from 56.8–85.5 in SVL.

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REDISCOVERY OF RHINELLA LEPTOSCELIS
FIGURE 1. (A–B) Holotype of Rhinella leptoscelis (BM 1907.5.7.32, SVL= 53.5 mm) from Santo Domingo, Carabaya,
Departamento Puno, Peru. (C–D) Neotype of R. veraguensis (BM 1947.2.21.23, SVL=41.0 mm; also lectotype of Bufo
ockendeni) from Marcapata Valley, Departmento Cusco, Peru.
Variation. We describe variation in qualitative characters, measurements, and color in life on the basis of
five newly collected specimens from the type locality (MHNC 5975–6, 5987–9; see associated data under
remarks section). No male specimens of R. leptoscelis have been found to date. The holotype and the five new
Peruvian specimens show few differences in qualitative characters (Fig. 1A–B and Fig. 2 A–G), mostly
related to skin texture and coloration. The largest adult female (MHNC 5976; Fig. 2B) has spiny skin texture.
The choanae and vomerine teeth were examined in an adult female with convoluted oviducts (MHNC 5975).
The choanae are large, round, and anterolateral; between them are the odonthophores, which are no more than
two tiny, almost indiscernible protuberances lacking vomerine teeth. Dorsal coloration is brownish-grey in
MHNC 5976 and brownish-orange in MHNC 5975, with a longitudinal middorsal pale yellow stripe (this
stripe is underlined with black in MHNC 5976). The dorsolateral row of tubercles is brownish-cream in
MHNC 5976 and intense orange in MHNC 5975. The flanks are pale grey with darker spots in MHNC 5976,
and orange and grey with tiny dark grey spots and white tubercles in MHNC 5975. In MHNC 5976, the lateral
surfaces of head are overall dark brownish-grey, with pale lips; dorsally, the eyelids and the interorbital region
are brownish-orange to cream, with two black spots posteriorly, and a longitudinal cream stripe; the occipital
region has two large yellow and bold black spots. In MHNC 5975, the head is overall orange, with a
middorsal longitudinal yellow stripe; the ventral regions have black, white, orange and grey flecks; the plantar
surfaces are dark brownish-grey, with yellowish-cream fringes and membranes. The hind limbs are dark grey
with two transversal cream to yellow narrow irregular stripes in MHNC 5976, and brownish-orange with
grayish-brown transversal bars in MHNC 5975. The forelegs are dark grey with some darker and paler spots
in MHNC 5976, and brownish-orange with grayish-brown transversal bars in MHNC 5975. Bolivian

PADIAL ET AL.60 · Zootaxa 2115 © 2009 Magnolia Press
specimens (see below) are also all females, with a light vertebral stripe, bordered by dark brown lines, and
cream lateral dorsum and flanks, contrasting with the darker mid-dorsum (Fig. 2F). Dark brown dorsal flecks
and markings are present in all Bolivian specimens. The throat is dark brown with some scattered cream spots
(Fig. 2G).
Remarks. Other specimens that we now consider to represent R. leptoscelis are those reported as
R. fissipes by Köhler (2000) for Bolivia. The identity of these Bolivian populations was reinvestigated and
some differences between the R. fissipes holotype and Bolivian specimens (i.e. tympanum condition)
mentioned by Köhler (2000), and the lack of distinct cranial crests in R. fissipes, seem to be reliable characters
to distinguish between both taxa. Consequently, R. leptoscelis, and not R. fissipes should now be listed for
Bolivia, although the presence of the latter in the country is very likely.
FIGURE 2. (A, D–E) Adult female of Rhinella leptoscelis from Santo Domingo de Carabaya, Departmento Puno, Peru
(MHNC 5975, SVL= 59.8 mm); (B) Adult female of R. leptoscelis from Santo Domingo de Carabaya, Departmento
Puno, Peru (MHNC 5976, SVL=85.5 mm); (C) Adult female of R. leptoscelis from Kimbiri River, 1550 m a.s.l.,
Departmento Cusco, Peru (MNCN 44406; SVL=65.7); (F–G) Adult female of R. leptoscelis from Chapare, Bolivia
(ZFMK 72670, SVL=65.6 mm).

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REDISCOVERY OF RHINELLA LEPTOSCELIS
Distribution. Since the discovery of Rhinella leptoscelis by Ockenden at the end of the Nineteenth
Century at Santo Domingo de Carabaya, Puno, Peru (2000 m a.s.l.) (Fig. 3), no additional specimens were
allocated to this species. One of the authors (JCC) rediscovered this species during an expedition to the type
locality and the surrounded area in November 2006. He collected two specimens (MHNC 5975–6) at Santo
Domingo de Carabaya, 1658 m a.s.l., and three specimens (MHNC 5987–9) at a lower altitude, 1400 m a.s.l.
(see Appendix). More recently, we collected additional specimens of R. leptoscelis (Fig. 2C) in Kimbiri River,
Vilcabamba Mountains, Cusco, Peru (Fig. 3; Appendix).
FIGURE 3. Map of the Central Andes depicting localities for Rhinella leptoscelis: (1) type locality at Santo Domingo de
Carabaya, 2000 m a.s.l, Departamento Puno, Peru; (2) Kimbiri River, 1350–1550 m a.s.l., Departmento Cusco, Peru; (3)
Chapare, 1300–1400 m a.s.l., Departmento Cochabamba, Bolivia; (4) Caranavi, Departamento La Paz, Bolivia.
Bolivian records of R. leptoscelis (see above) come from Chapare, 1300–1400 m a.s.l., Departamento
Cochabamba, and from the surroundings of Caranavi, Departamento La Paz, Bolivia (the latter specimens,
deposited at CBF, were erroneously reported as originating from Rurrenabaque, Departamento Beni, by
Köhler 2000).
Thus, Rhinella leptoscelis occurs in well-preserved humid montane forests from the Chapare region in
Bolivia to the Vilcabamba mountain range in southern Peru. All known specimens were found between
1300–1658 m. The species is known to occur in sympatry with R. veraguensis and R. fissipes at the type
locality, with R. inca in Vilcabamba, and with B. quechua, R. stanlaii, and R. veraguensis in the Bolivian
Chapare region.

PADIAL ET AL.62 · Zootaxa 2115 © 2009 Magnolia Press
Discussion
We have provided arguments for the recognition of R. leptoscelis as a valid species well distinguished from R.
veraguensis. But the assignation of Rhinella leptoscelis to the R. veraguensis group is tentative. The R.
veraguensis group is non-monophyletic, and some of its members are more closely related to members of the
R. m argaritifera group (Pramuk 2006; Chaparro et al. 2007). Rhinella leptoscelis shows a degree of
development in the orbytotympanic crests that resembles that of some members of the R. margaritifera group
(e.g. R. margaritifera, R. stanlaii). But the crests do not project above the tympanic area in R. leptoscelis, as is
typical in most members of the R. margaritifera group. Also, the morphological definitions of the R.
margaritifera and R. veraguensis groups provided by Duellman & Schulte (1992) were similar enough to
argue for a close relationship. Moreover, some of the species now assigned to the R. veraguensis group (R.
fissipes, R. inca, R. leptoscelis, and R. quechua) were former members of the R. margaritifera group
(Hoogmoed 1985). Thus, at least from the point of view of external characters, both groups are quite similar.
There is, however, an osteological synapomorphy (expansion of the posterior ramus of the pterygoid) shared
by some members of the R. margaritifera group (Pramuk 2006), which is not confirmed for current members
of the R. veraguensis group. Future molecular approaches will allow to improve our knowledge about the
phylogenetic relationships of all these species and to better define the limits, if any, between these species
groups.
Acknowledgments
JMP’s work has been partially financed by an Ernst Mayr Travel Grant in Animal Systematics (Museum of
Comparative Zoology, Harvard University), by the "European Commission's Research Infrastructure Action”
via the “SYNTHESYS Project" (GB-TAF-299) and by the “EU Marie Curie Mobility and Training
Programme (FP7). J.C. Chaparro’s fieldwork was funded by APECO and by a Koepcke grant. Collecting
permits were provided by the Instituto Nacional de Recursos Naturales (INRENA) to I. De la Riva. We are
grateful to the following persons for the loan of specimens, support and/or space provided at their institutions:
R. Bain, D. Frost, T. Grant and D. Kizirian, (AMNH), B. Clarke and M. Wilkinson (BM), J. Aparicio (CBF),
R. Aguayo and A. Muñoz (CBG), W. E. Duellman, J. Simmons and L. Trueb (KU), W. R. Heyer (USNM), J.
Hanken and J. Rosado (MCZ), O. Aguilar (MHNC), César Aguilar (MHNSM), J. E. González and T. García
(MNCN), L. González, A. Justiniano, R. Montaño, M. Suárez and R. Vespa (MNK), K. de Queiroz, W. R.
Heyer, R. McDiarmid and R. V. Wilson (USNM), and W. Böhme (ZFMK). We are grateful to S. Castroviejo
(EBC) and C. Vilà (EBD) for their help in the field in Vilcabamba. This work was partially funded by projects
REN/GLO 2001-1046 and CGL2005-03156 of the Spanish Ministry of Science and Innovation (I. De la Riva,
Principal Investigator).
Literature cited
Cei, J. M. (1968) Remarks on the geographical distribution and phyletic trends of South American toads. The Pearce-
Sellards Series, Texas Memorial Museum, 13, 1–21.
Cei, J. M. (1972) Bufo of South America. In: Blair, F.W. (Ed.), Evolution in the genus Bufo. University of Texas Press,
Austin, pp. 82–92.
Chaparro, J.C., Pramuk, J.B. & Gluesenkamp, A.G. (2007) A new species of arboreal Rhinella (Anura: Bufonidae) from
cloud forests of southeastern Peru. Herpetologica, 63, 203–212.
De la Riva, I., Köhler, J., Lötters, S. & Reichle, S. (2000) Ten years of research on Bolivian amphibians: updated
checklist, distribution, taxonomic problems, literature and iconography. Revista Española de Herpetología, 14,
19–164.
Duellman, W.E. & Schulte, 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, 162–172.

Zootaxa 2115 © 2009 Magnolia Press · 63
REDISCOVERY OF RHINELLA LEPTOSCELIS
Frost, D.R. (2009) Amphibian Species of the World: an Online Reference. Version 5.3 (12 February, 2009). American
Museum of Natural History, New York, USA. Electronic Database accessible at http://research.amnh.org/
herpetology/amphibia/index.php. (accessed 29 December 2008).
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing, A., Wilkinson, M.,
Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch,
J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural
History, 297, 1–370.
Gallardo, J.M. (1961) Three new toads from South America: Bufo manicorensis, Bufo spinulosus altiperuvianus and
Bufo quechua. Breviora, 141, 1–18.
Gorham, S.W. (1974) Checklist of world amphibians. New Brunswick Museum, St. Johns, Canada, 173 pp.
Harvey, M.B. & Smith, E.N. (1993) A new species of aquatic Bufo (Anura: Bufonidae) from cloud forest in the Serrania
de Siberia, Bolivia. Proceedings of the Biological Society of Washington, 106, 442–449.
Harvey, M.B. & Smith, E.N. (1994) A new species of Bufo (Anura: Bufonidae) from cloud forest in Bolivia.
Herpetologica, 50, 32–38.
Hayek, L.A., Heyer, R.W. & Gascon, C. (2001) Frog morphometrics: a cautionary tale. Alytes, 18, 153–177.
Hoogmoed, M. S. 1985. Bufonidae (part). In: Frost, D.R. (Ed.), Amphibian species of the world. A taxonomic and
geographical reference. Allen Press, Lawrence, pp. 27–77.
Hoogmoed, M.S. (1990) Biosystematics of South American Bufonidae, with special reference to the Bufo "typhonius"
group. In: Peters, G. & R. Hutterer (Eds.), Vertebrates in the tropics. Museum Alexander Koenig, Bonn, pp.
113–123.
Köhler, J. (2000) Amphibian diversity in Bolivia: a study with special reference to montane forest regions. Bonner
zoologische Monographien, 48, 1–243.
Lehr, E., Köhler, G., Aguilar, C. & Ponce, E. (2001) New Species of Bufo (Anura: Bufonidae) from Central Peru. Copeia,
2001, 216–223
Lehr, E., Pramuk, J.B. & Lundberg, M. (2005) A new species of Bufo (Anura: Bufonidae) from Andean Peru.
Herpetologica, 61, 308–318.
Padial, J.M., Reichle, S., McDiarmid, R. & De la Riva, I. (2006) A new species of arboreal Chaunus (Anura: Bufonidae)
from Madidi National Park, Bolivia. Zootaxa, 1278, 57–68.
Pramuk, J.B. (2006) Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence.
Zoological Journal of the Linnean Society, 146, 407–452.
Savage, J.M. (1969) Clarification of the status of the toad, Bufo veraguensis O. Schmidt, 1857. Copeia, 1, 178–179.
Vellard, J. (1959) Estudios sobre batracios andinos. V. El género Bufo. Memorias del Museo de Historial Natural "Javier
Prado", 8, 1–48.

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Appendix
Specimens examined
Rhinella amboroensis. BOLIVIA: Departamento Cochabamba: 12.7 km by road E of El Empalme along road to Khara
Huasi, MNK-A 953 (holotype).
Rhinella fissipes. PERU: Santo Domingo, Carabaya, S.E. Peru, 6000 feet, BM 1947.2.20.64 (holotype).
Rhinella inca. PERU: Departamento Cusco: Huadquinia, USNM 49557 (holotype); Kimbiri River, 1100 m a.s.l.,
12º35'26.5"S, 73º41'36.8"W, MNCN 44405.
Rhinella justinianoi. BOLIVIA: Departamento Cochabamba: Karahuasi, 1800 m a.s.l., ZFMK 72657; Old Chapare road,
1650 m a.s.l., ZFMK 72600–02; Old Chapare road 2250 m a.s.l., ZFMK 72621; Saquisacha, CBG 168–176;
Departamento Santa Cruz: El Chapé, MNK-A950 (holotype).
Rhinella leptoscelis. BOLIVIA: Departamento Cochabamba: Chapare 1300-1400 m a.s.l., ZFMK 66985, 72668–71,
80035–36; PERU: Departamento Puno: Santo Domingo, Carabaya, S.E. Peru, 6500 feet, BM 1907.5.7.32; Santo
Domingo de Carabaya 1658 m a.s.l., 13º49’59.6’’S, 69º38’31.8’’W, MHNC 5975–76; Santo Domingo de Carabaya,
1400 m a.s.l., 13º46’14.3’’S, 69º37’28.7’’W, MHNC 5987–89; Departamento Cusco: Kimbiri River 1350-1550 m
a.s.l., 12º34'12.4"S, 73º39'17.6"W, MHNC 6803, 6812, 6817, MNCN 44404, 44406.
Rhinella manu. PERU: Departamento Cusco: Tres Cruces, approx. 134 km NE of Cusco, 2750 m a.s.l., MHNC 3005
(holotype), MHNC 3003–4, 3006–11, MHNSM 24883 (paratypes).
Rhinella quechua. BOLIVIA: Departamento Cochabamba: Incachaca (type locality), 2300 m a.s.l., ZFMK 66939–41;
Old Chapare Road, 2250 m a.s.l., EBD 30249–30252, ZFMK 72622; Sehuencas, 2200 m a.s.l., CBG 109–127,
ZFMK 60255–74, 60276–82, 66835–36.
Rhinella stanlaii. BOLIVIA: Departamento Cochabamba: road Villa Tunari-Cochabamba, 1850 m a.s.l., ZFMK 60464
(paratype); road to San Onofre, 1900 m a.s.l., CBF 3346 (holotype); Old Chapare road, 1400 m a.s.l., ZFMK 67096
(paratype); Departamento Santa Cruz: La Hoyada, 1700 m a.s.l., ZSM 144/1999 (paratype).
Rhinella tacana. BOLIVIA: Departamento La Paz: Arroyo Huacataya, path from San José de Uchupiamonas to Apolo,
Serranía Eslabón, Madidi National Park, MNCN 42073, MNK-A-7194 (paratypes); Huairuro path from San José de
Uchupiamonas to Apolo, Serranía Eslabón, Madidi National Park, MNK-A 7188 (holotype), MNK-A 7187, MNCN
42072 (paratypes).
Rhinella veraguensis (=Bufo ockendeni). BOLIVIA: Departamento Beni: Río Yucumo, 5 km upstream from Yucumo,
MNCN 43836-37; Departamento Cochabamba: Campamento Los Guácharos, Carrasco National Park, MNCN
42633, 43017; Charuplaya, BM 1947.2.21.26-27 (paralectotypes); Karahuasi, CBG 226–236, ZFMK 72658; Old
Chapare road, 1250 m a.s.l., ZFMK 72555–58; La Siberia, road to Locotal, MNCN 43369; Old Chapare road,
1300–1500 m a.s.l., ZFMK 72574–75; Old Chapare road, 1650 m a.s.l., ZFMK 72590–92; Quebrada on the road
Cochabamba-Villa Tunari, MNCN 43379; road Los Guácharos-El Palmar, 8 km from Los Guácharos, MNCN
43032; 120 km from Cochabamba on Cochabamba-Villa Tunari road, MNCN 43384; Departamento La Paz: Arroyo
Huacataya, path between San José de Uchupiamonas and Apolo, MNCN 43414; Puesto Guardaparques-Arroyo
Huabudahaida, Madidi National Park, MNCN 43011; Serranía de Bella Vista, MNK-A7270-71, MNCN 41991;
Valle de Zongo, MNCN 43413; Departamento Santa Cruz: El Empalme, Serranía de la Siberia, MNCN 43736; El
Fuerte, Samaipata, ZFMK 62832-33, 66884; 29 km SE of Guadalupe, 1600 m a.s.l., ZFMK 66850–51; La Yunga,
2300 m a.s.l., MNCN 43370–71, ZFMK 66880; 45 km W of Río Seco, ZFMK 67077–78; 120 km from Santa Cruz
to Cochabamba, MNCN 43380–81; PERU: Departamento Cusco: Marcapata Valley, BM 1947.2.21.23 (lectotype).