Zootaxa 1278: 57-68 (2006) ISSN 1175-5326 (print edition) 
www.mapress.com/zootaxa/ 7C\C\rI1 A "Y" \ {*\^HS?\ 
Copyright ? 2006 Magnolia Press ISSN , ^5.5334 (online edition) 
A new species of arboreal toad (Anura: Bufonidae: Chaunus) from 
Madidi National Park, Bolivia 
JOSE M. PADIAL1, STEFFEN REICHLE2, ROY McDIARMID3 & IGNACIO DE LA 
RIVA1* 
'Department of Biodiversity and Evolutionary Biology, Museo Nacional de Ciencias Naturales, CSIC. C/ Jose 
Gutierrez Abascal 2, 28006 Madrid, Spain 
2Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany 
3USGS Patuxent Wildlife Research Center, Smithsonian Institution, National Museum of Natural History, PO 
Box 37012, Washington, DC 20013-7012 USA 
'^Corresponding author: iriva@mncn.csic.es 
Abstract 
A new arboreal species of the Chaunus veraguensis group is described for the humid montane 
forest of Madidi National Park, in northern Bolivia. The new species differs from other species in 
the group by the combination small size, long and slender extremities, webbed hands, conspicuous 
tympanic membrane, well developed parotoid glands, absence of large glands on dorsum and 
extremities, nuptial excrescences of males composed of pungent spines on dorsal surface of thumb, 
greenish-brown coloration on dorsum with red warts in life, and green iris. It is only known from 
two nearby localities in the Serrania Eslabon, Department La Paz. An operational key for species in 
the C. veraguensis group is provided. 
Key words: Anura, Bufonidae, Chaunus veraguensis group, Bolivia, Andes, humid montane 
forest, new species 
Resumen 
Se describe una nueva especie arboricola del grupo Chaunus veraguensis de los bosques montanos 
humedos andinos del Parque Nacional Madidi, norte de Bolivia. La nueva especie se diferencia de 
otras especies del grupo por su pequefio tamano, por poseer las extremidades largas y delgadas, las 
plantas de las manos y de los pies palmeadas, la membrana timpanica visible y conspicua, y las 
glandulas parotidas bien desarrolladas y redondeadas, por la ausencia de otro tipo de grandes 
glandulas en dorso o extremidades, por las espinas nupciales negras de la region dorsal del pulgar 
de los machos, por su coloracion verdosa en vida y por su iris verde. Solo es conocida de dos 
localidades cercanas en la Serrania Eslabon, en el Departmento de La Paz. Se proporciona una 
clave de identification para las especies del grupo C. veraguensis. 
Accepted by M. Vences: 23 Jun 2006; published: 3 Aug. 2006 57 
ZOOTAXA Palabras clave: Anura, Bufonidae, grupo Chaunus veraguensis, Bolivia, Andes, bosque humedo 
(XVTft) de montana, new species 
Introduction 
The Andean foothills of northern Bolivia, mainly protected by the Madidi National Park 
and the Area Nacional de Manejo Integrado Apolobamba, conform one of the hottest 
biodiversity corners on earth (Remsem and Parker 1995). However, amphibians have still 
not received much attention and the scarce information comes from a few expeditions. 
Only recently, this area has attracted the interest of herpetologists, which resulted in some 
interesting discoveries including the description of new species (Cortez 2005; De la Riva 
et al. 2005a, b; Padial et al. 2005). During a short survey of the Serrania Eslabon in the 
Madidi National Park, the senior author collected five specimens of a completely arboreal 
toad similar to species of the Chaunus veraguensis group that did not correspond to any 
known species from northern Bolivia or Southern Peru. Comparisons of this species with 
museum specimens and original descriptions of other Bolivian and Peruvian species from 
various groups, convinced us that our specimens belong to an undescribed species. The 
aim of this article is to describe and name this new species. Moreover, we provide an 
operational key for the species currently included in the C. veraguensis group. 
Material and methods 
Specimens were fixed in 10% formalin and preserved in 70% ethanol. For morphological 
and colour 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 calliper to the nearest 0.01mm, but following Hayek et al. (2001), for 
avoiding pseudo precision, we rounded all measurements to only one decimal. Webbing 
formulae follow Savage and Heyer (1967) as modified by Myers and Duellman (1982) and 
Savage and Heyer (1997). 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 narines, IND; distance between the anterior 
margin of the eyes (eye-eye distance), EE; tympanic membrane height, TYH; tympanic 
membrane length, TYL; forearm length (from posterior margin of thenar tubercle to 
elbow), FA; tibia length, TL; thigh length, TH; foot length (from proximal border of inner 
metatarsal tubercle to tip of fourth toe), FL. Color characteristics were noted in life. 
Museum abbreviations refer to: Centra de Biodiversidad y Genetica, Cochabamba, Bolivia 
(CBG); Coleccion Boliviana de Fauna, La Paz, Bolivia (CBF); Museo de Historia Natural 
Noel   Kempff  Mercado,   Santa   Cruz   de   la   Sierra,   Bolivia   (MNK-A   [Amphibian 
58 ? 2006 Magnolia Press PADIAL ET AL. 
Collection]); Estacion Biologica de Doiiana, Sevilla, Spain (EBD); Museo Nacional de ZOOTAXA 
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). 
Results 
Chaunus tacana sp. nov. 
Fig 1, 3-4 
Holotype 
MNK-A 7188, an adult female from Huairuro (14?19'28.2" S, 68?05'36.1" W), path 
from San Jose de Uchupiamonas to Apolo, Serrania Eslabon, Madidi National Park, 
Province Franz Tamayo, Department La Paz, Bolivia (Fig. 2), collected the night of 16-17 
December 2003 by Jose M. Padial (field number JMP 556). 
Paratypes 
MNK-A 7187 (field number JMP 555), an adult male (Fig. 3) and MNCN 42072 (field 
number JMP-558), an immature female, both collected with the holotype; an adult male 
MNCN 42073 (field number JMP-574) and a juvenile MNK-A-7194 (JMP575) collected 
at Arroyo Huacataya (14?20'12.1" S, 68?5'57.3" W), path from San Jose de Uchupiamonas 
to Apolo, Serrania Eslabon, Madidi National Park, Department La Paz, Province Franz 
Tamayo, Bolivia, collected the night of 17-18 December 2003 by Jose M. Padial, Pere 
Comas and Pedro Macuapa. 
Etymology 
The specific name is a substantive in apposition that refers to the Bolivian indigenous 
group Tacana, who inhabit the lowlands and foothills of the Madidi National Park. The 
Tacana people have successfully demonstrated (through management of the Chalalan Eco- 
lodge) that community development is compatible with conservation of their cultural and 
natural heritage. 
Diagnosis 
A small species of Chaunus with long and slender extremities, tentatively assigned to 
the C. veraguensis group sensu Duellman and Schulte (1992) based on external characters. 
The new species is distinguished from other putatively related species by the following 
combination of characters: (1) canthus rostralis sharp, orbitotympanic and postorbital crest 
weak; (2) tympanum distinct, oval; (3) parotoid glands large, round, not contacting the 
tympanic membrane; (4) numerous scattered small, round warts on dorsal surfaces of 
body; (5) extremities moderately long and slender; (6) a dorsolateral row of round, 
ANEW CHAUNUS ? 2006 Magnolia Press 59 
(127?) 
ZOOTAXA enlarged, and elevated warts; (7) tarsal fold absent; (8) webbing on hands and feet fleshy 
(Figure 3); (9) first finger shorter than second (figure 3); (10) males with vocal slits; (11) 
small keratinous spines on dorsal surfaces of thumb; (12) iris green in life. 
The most similar species is one that is being described by J. A. Chaparro, J. Pramuk 
and A. Gluesenkamp from Department Cusco, Peru. We examined specimens of this 
undescribed taxon and compared them with C. tacana. Both species are similar in habitus, 
both are arboreal and inhabit cloud forests and humid montane forests. Nevertheless, C. 
tacana differs from the species from Cusco as follows (characteristics of the latter in 
parentheses): fingers thin and long (thick and short); webbing of hands and feet less 
developed, almost absent between Finger I and II (well developed between finger I and II); 
tarsal fold absent (present); parotoid glands proportionally smaller; parotoid glands and 
tympanic membrane not in contact (in contact); vocal slits present in males (absent); 
dorsal coloration in life dorsally pale greenish-brown to brownish-green with brown 
stripes or spots and red warts (homogeneously black); ventral coloration whitish-cream 
with light brown reticulation (red with dark blotches); iris green (reddish-orange). 
Chaunus tacana can be distinguished from another similar and arboreal species, C. 
arborescandens, mainly by the visible tympanic membrane (absent in the latter), and the 
concave internarial area (flat). Chaunus tacana has visible tympanic membrane while C. 
amboroensis, C. justinianoi, C. quechua, and C. veraguensis lack visible tympanic 
membrane. C. fissipes may have a visible tympanum although barely evident. 
Nevertheless, C. fissipes is larger [mean adult females SVL= 68.1 (range= 65.8-71.9, 
n=8); see Kohler (2000)] and has two prominent dorsolateral rows of tubercles. Moreover, 
C. fissipes has the first finger longer than second, the parotoid glands are slightly elevated 
and not protruding laterally, and it presents serrate foot webbing. The new species can be 
distinguished from C. multiverrucosus and C. chavin by the nuptial spines on the first 
finger in breeding males, and by the lack of enlarged glands on dorsum and extremities. 
Chaunus tacana differs from C. inca by having the first finger shorter than second. The 
new species differs from C. nesiotes by having pronounced parotoid glands (low and 
diffuse in C. nesiotes) and a row of enlarged dorsolateral warts (absent). C. tacana differs 
from C. rumbolli by the long and slender extremities, inconspicuous cranial crests and first 
finger longer than second (shorter in C. rumbolli). 
Description of the holotype 
A small Chaunus with long and robust body (for measurements see table 1); head 
small, slightly broader than long, its width 30% of SVL, length 30% of SVL; snout short, 
subacuminate in dorsal view, acute in profile, with a vertical keel on the tip; cranial crests 
absent; skin of head not coosified with underlying cranial bones; internarial area concave; 
narines not protuberant, very small, oriented laterally; canhus rostralis sharp, concave in 
dorsal view; lips flat; eye-nostril distance equal to eye length; tympanic membrane visible, 
conspicuous, oval, surrounded by granules, 70% of eye length; tympanic annulus thin, 
overlapped with surrounding granules. Forelimb long and slender; hand broad, with short 
60 ? 2006 Magnolia Press PADIAL ETAL. 
fingers; relative length of fingers I<II<IV<III; basal webbing fleshy, extending as a fringe 
on lateral edges of fingers; webbing formula i12nbasalIIIbasalIV; tips of digits round, not 
expanded; ulnar region granular with a row of enlarged tubercles; palmar tubercle round 
and low; prepollical tubercle irregular to ovoid, diffuse; subarticular tubercles ovoid to 
round, low, diffuse, inconspicuous; supernumerary tubercles round, diffuse, smaller than 
subarticular. Hind limbs and feet long; tibia length 40% of SVL; foot length 40% of SVL; 
no tarsal fold; outer metatarsal tubercle ovate, prominent, 1/3 the size of inner; inner 
metatarsal tubercle large, prominent, ovate; relative length of toes 1<2=5<3<4; webbing 
fleshy, almost complete, but last phalange free, only basal between toes 4 and 5; webbing 
formula l1/21ll1/21in1/2"3IV3"2V; tip of toes rounded; subarticular tubercles low, round to 
ovate, diffuse, larger than supernumerary tubercles; supernumerary tubercles abundant, 
small, irregular or rounded. 
ZOOTAXA 
(127?) 
TABLE 1. Measurements and proportions of the adult type specimens of Chaunus tacana sp. nov. 
(M, male; F, female; for other abbreviations see Material and Methods). 
NKA-A 7188 (F)      NKA-A 7187 (M) MNCN 42073 (M) 
SVL 
HL 
HW 
EW 
ED 
EN 
IND 
EE 
TYH 
TYL 
FA 
TL 
TH 
FL 
HW/HL 
HW/SVL 
HL/SVL 
EN/ED 
EE/ED 
TYL/ED 
TL/SVL 
FL/SVL 
34.2 30.6 
9.9 9.2 
10.8 9.7 
2.8 2.5 
3.1 3.3 
3.0 2.9 
2.4 1.9 
5.0 4.8 
2.7 2.2 
2.1 2.0 
8.3 7.6 
12.3 12.7 
12.9 11.4 
14.2 14.2 
1.1 1.1 
0.3 0.3 
0.3 0.3 
1.0 0.9 
1.6 1.5 
0.7 0.6 
0.4 0.4 
0.4 0.5 
30.7 
9.1 
9.4 
2.8 
3.5 
2.7 
1.9 
4.5 
2.2 
2.0 
6.8 
11.6 
11.7 
12.6 
1.0 
0.3 
0.3 
0.8 
1.3 
0.6 
0.4 
0.4 
ANEW CHAUNUS ? 2006 Magnolia Press 61 
ZOOTAXA Skin on dorsum of head, body, and limbs bearing abundant small, flat granules of 
irregular size and low rounded warts, larger than granules; enlarged granules forming an 
irregular row that resembles a dorsolateral fold; parotoid gland round, large, 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 crests composed of 
granules; enlarged glands absent on limbs. Skin on throat and other ventral surfaces 
granular. Anal opening slightly protuberant, directed posteroventrally at upper level of 
thighs. 
One choanae small and round, placed anterolaterally, while the other is concealed by 
the palatal shelf of the maxillary arch; vomerine odontophores absent; tongue cordiform, 
free behind for about one-third its length. 
Color in preservative: Dorsum dark grey, with enlarged light brown warts; interocular, 
scapular, and central areas of dorsum with a bold black irregular x-shaped stripe; granules 
forming a dorsolateral row cream; flanks scarcely lighter than dorsum; tympanic 
membrane dark greyish-brown; head dark greyish-brown, lighter in lower parts; forelegs 
barred and hind legs greyish-brown with dark brown, almost black, transverse bars; toes 
barred with dark brown; ventral regions irregularly mottled with brown and cream, lighter 
at the level of groin; plantar surfaces cream with some brown spots. 
Color in life: dorsal coloration pale brown to greenish-brown; scapular and central 
areas of dorsum with a dark brown black irregular x-shaped stripe with greenish-brown 
tonalities; granules forming the dorsolateral rows brown or red; tympanic membrane 
brown; head brown with some red granules and small greenish-brown spots on interorbital 
region; extremities brown, barred with dark brown to greenish-brown; toes barred with 
dark brown; ventral coloration whitish-cream with light brown reticulation; iris green, 
pupil black. 
Variation 
Little variation is observed. There is little sexual dimorphism in size. Males present 
nuptial excrescences in form of a single group of black (in life) or white (in alcohol) 
keratinized spines on the dorsal surface of each thumb. Male specimens present vocal slits 
lateral to tongue. Males seem to have shorter snout relative to eye length (80%) than 
females (100%), and also narrower snout (EN 160% of EE distance in females versus 
130% in males). In one male (MNCN 42073) both choanae are visible, they are 
anterolateral and are separated by a distance equal to six times the diameter of a choanae. 
Most spicules and granules can be concentrated on neck and head, while the flanks have 
larger granules and warts but density of granules is lower. In MNCN 42073 the parotoid 
gland is slightly ovate, displaced to the lateral part of the head, where it reaches the level 
of the lower margin of the tympanic membrane. In MNCN 42073 the foot webbing 
formula differs slightly from that of the holotype: i^ii^m^iv^y. Juveniles are 
identical to adults in general appearance. Dorsal pattern varies slightly in the intensity and 
62 ? 2006 Magnolia Press PADIAL ETAL. 
form of the stripes. Some individuals have more red warts. SVL of two juveniles: 25.3 
(MNCN 42072, female) and 21.8 (NKA-A7194, unsexed). NKA-A 7187 lack dorsal 
pattern and is darker than any other specimen in alcohol (Fig. 3). 
ZOOTAXA 
(127?) 
FIGURE 1. Adult female holotype of Chaunus tacana sp. nov. from Serrania Eslabon, Madidi 
National Park, Bolivia (MNK-A 7188) 
Distribution and natural history 
This species is known only from two close localities in Serrania Eslabon (Fig. 2), at 
about 1500 m a.s.l. The habitat is composed by primary humid montane forest with high 
abundance of arboreal ferns, bromeliads and other epiphytes. Potentially flooded flat 
surfaces are almost inexistent but there are some small streams. C. tacana is a nocturnal 
species that can be found active on leaves of bushes or on the trunk of trees from 1-4 m 
height. It climbs graciously vertical trunks covered by moss. As C. tacana, C. veraguensis 
may also show greenish dorsal coloration and has been reported to climb in trees (Kohler 
2000). This mimetic coloration could be related to their arboreal habits. The adult female 
presented convoluted oviducts and small white ova. The call, tadpole and reproductive 
mode are unknown. Other anuran species that occur in the area are Atelopus tricolor, 
Epipedobates bolivianus, Hyalinobatrachium bergeri, Hyloscirtus armatus, Hypsiboas 
balzani, Eleutherodactylus danae and E. madidi. 
) M\*\ 1000 m 
| 3000 m 
t 
%   ^ 
ft 
&4 v 
-s^.              N 
>      La Paz    ^ 
Jo V; 5& r^ w._ Sanla Cruzj 
100 km 1 *   (S n 
FIGURE 2. Map of Bolivia with a bold square indicating the type locality of Chaunus tacana sp. 
nov. 
ANEW CHAUNUS ? 2006 Magnolia Press 63 
ZOOTAXA 
FIGURE 3. Dorsal and ventral view of an adult male of Chaunus tacana sp. nov. from Serranfa 
Eslabon, Madidi National Park, Bolivia (MNK-A 7187). 
FIGURE 4. Dorsal view of and adult female of Chaunus sp. (left) from Department Cusco, Peru 
and the holotype (adult female) of Chaunus tacana sp. nov (MNK-A 7188). 
Discussion 
We follow the recent taxonomic rearrangements proposed by Frost et al. (2006) for the 
genus Bufo. They recognized the genus Chaunus (Wagler, 1828) for a ""...major clade of 
predominantly Neotropical "Bufo" (exluding Rhinella)..."" including the B. veraguensis 
group. Hence, the new species is herein assigned to the Chaunus veraguensis species 
group (former Bufo veraguensis species group) because of its external similarity to some 
of the species assigned to this group by Duellman and Schulte (1992). Thus, the Chaunus 
veraguensis group is currently composed of 12 species; apart from C. tacana these are: C. 
amboroensis (Harvey and Smith), C. arborescandens (Duellman and Schulte), C. chavin 
(Lehr, Kohler, Aguilar and Ponce), C. fissipes (Boulenger), C. inca (Stejneger), C. 
justinianoi (Harvey and Smith), C. nesiotes (Duellman and Toft), C. quechua (Gallardo), 
C. multiverrucosus (Lehr, Pramuk and Lundberg), C. rumbolli (Carrizo) and B. 
veraguensis (Schmidt). An additional species of the group is currently being described 
64 ? 2006 Magnolia Press PADIAL ETAL. 
from Department Cusco, Peru (J. C. Chaparro, pers. comm.). Seven described species ZOOTAXA 
occur in Bolivia (C amboroensis, C. fissipes, C. justinianoi, C. quechua, C. rumbolli, C. 
veraguensis and C. tacana) and also seven in Peru (C. arborescandens, C. chavin, C. 
fissipes, C. inca, C. nesiotes, C. veraguensis, C. multiverrucosus). Moreover, the presence 
of C. tacana is predicted for southern Peru. Nevertheless, De la Riva et al. (2000) already 
pointed out that some species in this group remain to be described, whereas the taxonomic 
status of others is doubtful. Hence, the real diversity of this group is still unknown. This 
assumption is supported by the high number of new species assigned to the C. veraguensis 
group described recently (Duellman and Schulte 1992; Harvey and Smith 1993; Harvey 
and Smith 1994; Lehr et al. 2001; Lehr et al. 2005). Moreover, infrageneric divisions, as 
species group in this case, are mostly based on similarities of external characters and no 
synapomorphies have been proposed so far. Thus, assignation to species groups is 
tentative. Nevertheless, the subdivision of genera in species groups helps to work with 
more manageable units, and there is the hope that some of them actually represent 
evolutionary units. Indeed, there are already attempts to clarify these phylogenetic 
relationships (e. g. Pramuk 2006). However, due to the number of new species of Chaunus 
that have been described recently after the exploration of new regions, it is easy to 
hypothesize that the number of species will increase and, therefore, the hypotheses on 
relationships among some species and groups would probably change in the near future. 
Operational key to the species of the Chaunus veraguensis group 
1. Tympanic membrane visible 2 
Tympanic membrane not discernible 8 
2. Large glands on forearm and hind legs 3 
No such glands 4 
3. Dorsum with many, large, elevated, conical glands, all with keratinous tips, skin spiny 
C. multiverrucosus 
Dorsum with few large, slightly elevated, ovoid glands, glands with keratinous tips 
restricted to head and neck, skin smooth C. chavin 
4. Parotoid glands diffuse C. nesiotes 
Parotoid glands evident 5 
5. First finger shorter than second 6 
First finger longer than second  C. inca 
6. Extremities robust, cranial crests evident  C. rumbolli 
Extremities slender, cranial crest scarcely evident or absent, males with nuptial spines 
7 
7. Dorsal coloration greenish, ventral brown and white, vocal slits present, iris green  
C. tacana 
Dorsal coloration black, ventral red, vocal slits absent, iris orange Chaunus sp. (Cusco) 
ANEW CHAUNUS ? 2006 Magnolia Press 65 
(127?) 
ZOOTAXA 8.   Cranial crests absent 9 
Cranial crests present 11 
9. Foot webbing complete C. amboroensis 
Foot webbing rudimentary 10 
10. Foot webbing fleshy C. arborescandens 
Foot webbing serrated C.fissipes 
11. First finger shorter than second C. justinianoi 
First finger longer than second 12 
12. Supraorbital crest absent; tarsal fold absent; parotoid gland elongate C. quechua 
Supraorbital crest present; tarsal fold consisting of a single row of granules; parotoid 
gland subtriangular C. veraguensis 
Acknowledgments 
For the help provided during many years, we are grateful to the personnel at the MNK, 
especially to M. Suarez, R. Vespa, A. Justiniano, and R. Montano. R. Aguayo (CBG), C. 
Aguilar and J. Cordova (MHNSM), J. Aparicio (CBF), J. Cabot (EBD), W. Bohme and J. 
Kohler (ZFMK), B. Clarke (BM) and W. R. Heyer (USNM), provided working facilities 
and specimens for study. We thank to P. Comas, D. Embert, and P. Macuapa for their 
company during the fieldwork. We are indebted to Juan Carlos Chaparro, who kindly 
allowed JMP and ID1R to examine the specimen of his new species under description. The 
director of the Madidi National Park, O. Loayza, kindly accepted our research project. We 
are extremely grateful to the people of San Jose de Uchupiamonas and to all the personnel 
of the Chalalan Ecolodge, for their support, help, and friendship, and we are especially 
indebted to its manager G. Mamani, for allowing us to use the outstanding Chalalan 
facilities as our base camp. Padial's research was financed by a grant of the Mutis 
programme of the MAE-AECI (Spain), his trip to visit the museums in USA was funded 
by an Ernst Mayr Travel Grant in Animal Systematics (Museum of Comparative Zoology, 
Harvard University) and his trip to BM was founded by a Synthesys grant of the EU. This 
work was partially funded by projects REN/GLO 2001-1046 and CGL2005-03156 of the 
Spanish Ministry of Education and Science (I. De la Riva, Principal Investigator) and by a 
project of the AECI (Spanish Agency of International Cooperation) (I. De la Riva, 
Principal Investigator) to inventory Madidi's herpetofauna. 
Literature cited 
Cortez, C. (2005) Herpetofauna de la zona norte del Parque Nacional y Area Natural de Manejo 
Integrado Madidi (PNANMI-Madidi). Ecologia en Bolivia, 40 (2), 10-26. 
De la Riva, I., Kohler, J., Lotters, S. & Reichle, S. (2000) Ten years of research on Bolivian 
66 ? 2006 Magnolia Press PADIAL ETAL. 
amphibians: updated checklist, distribution, taxonomic problems, literature and iconography. ZOOTAXA 
Revista Espanola de Herpetologia, 14, 19-164. (X27o) 
De la Riva, I., Rfos, N. & Aparicio, J. (2005a) A new species of Bufo (Anura: Bufonidae) from the 
Andes of Bolivia. Herpetologica, 61 (3), 280-286. 
De la Riva, I., Ribs, N. & Aparicio, J. (2005b) New Species of Telmatobius (Anura: Leptodactyl- 
idae) from Humid Paramo of Peru and Bolivia. Journal of Herpetology, 39 (3), 409^-16. 
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. 
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sa, 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. 
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, LA., Heyer, R.W. & Gascon, C. (2001) Frog morphometrics: a cautionary tale. Alytes, 18, 
153-177. 
Kohler, J. (2000) Amphibian diversity in Bolivia: a study with special reference to montane forest 
regions. Bonner zoologische Monographien, 48, 1-243. 
Lehr, E., Kohler, 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., Gonzales, L. & De la Riva, I. (2005) A new species of the Eleutherodactylus discoida- 
lis Group (Anura: Leptodactylidae) from Andean Humid Montane Forest of Bolivia. Herpeto- 
logica, 61 (3), 318-325. 
Myers, C.W. & Duellman, W.E. (1982) A new species of Hyla from Cerro Colorado, and other tree 
frog records and geographical notes from Western Panama. American Museum Novitates, 
2752, 1-32. 
Pramuk, J.B. (2006) Phylogeny of South American Bufo (Anura: Bufonidae) inferred from com- 
bined evidence. Zoological Journal of the Linnean Society, 146, 407-452. 
Remsen, J.V. & Parker, T.A. (1995) Bolivia has the opportunity to create the planet's richest park of 
terrestrial biota. Bird Conservation International, 5, 181-199. 
Savage, J.M. & Heyer, WR. (1967) Variation and distribution in the tree-frog genus Phyllomedusa 
in Costa Rica, Central America. Beitrge zur Neotropischen Fauna, 5 (2), 111-131. 
Savage, J.M. & Heyer, W.R. (1997). Digital webbing formulae for anurans: A refinement. Herpeto- 
logical Review, 28 (3), 131. 
ANEW CHAUNUS ? 2006 Magnolia Press 67 
ZOOTAXA Appendix 
Specimens examined 
Bufo leptoscelis (=C. veraguensis). Peru: Santo Domingo, Carabaya, Department Puno 
(Holotype: BM 1907.5.7.32). 
Bufo ockendeni (=C. veraguensis). Peru: Marcapata Valley, Department Puno (Lectotype: BM 
1907.2.21.23). Bolivia: Charuplaya (Paralectotypes: BM 1907.2.21.26-27). 
Chaunus amboroensis. Bolivia: 12.7 km by road E of El Empalme along road to Khara Huasi, 
Department Cochabamba (Holotype: MNK-A 953). 
Chaunus fissipes. Bolivia: Old Chapare Road, Department Cochabamba (ZFMK 66985). 
Chaunus justinianoi. Bolivia: El Chape, Department Santa Cruz (Holotype: MNK-A950); Old 
Chapare road, Department Cochabamba (ZFMK 72600-02); Saquisacha, Department Cochabamba 
(CBG 168-176) 
Chaunus quechua. Bolivia: Road from Cochabamba to Villa Tunari, Department Cochabamba 
(EBD 30249-30252); Sehuencas, Department Cochabamba (CBG 109-127; ZFMK 60255-74). 
Chaunus veraguensis. Bolivia: Samaipata, Department Santa Cruz (ZFMK 62832-3, 66884); 
45 km W of Rio Seco, Department Santa Cruz (ZFMK 67077-8); Serrana de Bella Vista, 
Department La Paz (MNK-A7270-1); Khara Huasi, Department Cochabamba (CBG 226-236). 
68 ? 2006 Magnolia Press PADIAL ETAL.