Taxonomic Reassessment of the Arboreal Toad Genus Pedostibes Gu¨ nther 1876
(Anura: Bufonidae) and Some Allied Oriental Bufonid Genera
AND A.A. THASUN AMARASINGHE
Wildlife Institute of India, P.O. Box 18, Chandrabani, Dehradun 248001, India
Research Center for Climate Change, University of Indonesia, Gd. PAU Lt. 8.5, Kampus UI, Depok 16424, Indonesia
ABSTRACT: We reassessed the taxonomic status of an Asian genus of arboreal bufonids, Pedostibes, based on examination of preserved material of
the two species currently attributed to this genus. Analysis of their morphological, morphometric, and geographic distribution data revealed that
Pedostibes tuberculosus, the type species of this genus from the Western Ghats, southwestern India, is morphologically distinct from the
geographically separated member, P. kempi, which is distributed in northeastern India. Hence, the generic nomen Pedostibes is restricted to the type
species, rendering it a monotypic genus from the Western Ghats of peninsular India. A re-examination and detailed comparisons of the types of P.
kempi with other bufonid genera revealed morphological similarities with another geographically proximate toad, Bufoides meghalayanus, from
northeastern India. Hence, this taxon is formally transferred herein to Bufoides with a redescription. The composition of the recently described
Southeast Asian toad genus Rentapia is reevaluated and the name-bearing type specimens of the currently ascribed taxa are redescribed. A detailed
examination of the types of Rentapia everetti and R. rugosa revealed morphological congruence coupled with geographic sympatry. Hence, the latter
nomen is synonymized with R. everetti in accordance with the International Code of Zoological Nomenclature principle of priority.
Key words: Arboreal anuran; Morphometric analyses; Northeastern India; Western Ghats
THE ASIAN genus of arboreal bufonids, Pedostibes
Gu¨ nther 1876, previously show n to be polyphyletic
(Bocxlaer et al. 2009; Pyron and Wiens 2011; Ron et al.
2015), has recently been revised by Chan et al. (2016) with
the recognition of a new Sundaic genus, Rentapia Chan,
Grismer, Zachariah et al. 2016. Although this step has
contributed toward providing a taxonomic solution to the
problem of polyphyly in Pedostibes to a certain extent, the
classification and rearrangement therein has not provided
complete taxonomic clarity to Pedostibes. Not having
examined any specimens of the taxa Pedostibes kempi and
P. everetti, which were ascribed to Pedostibes earlier, Chan
et al. (2016) made an explicit remark on their uncertain
generic status, and provisionally allocated them to the
genera Pedostibes and Rentapia respectively. Although
Chan et al. (2016) provided evidence for differentiation of
the “Indian” and Sundaic lineages, they did not provide
a comprehensive description of the included taxa. Thus,
even after the rearrangement by Chan et al. (2016), the
genus Pedostibes still shows a disjunct pattern in geographic
distribution with two representative species, P. tuberculosus
Gu¨ nther 1876 and P. kempi (Boulenger 1919) occurring in
different biogeographic regions—the Western Ghats and
the Garo Hills in the eastern Himalayas (both within the
political boundary of India). In order to improve the
taxonomic resolution of this group, we present results based
on morphological examination of the name-bearing type
specimens for all of the relevant taxa from these regions and
provide detailed redescriptions of the species along with
some justified taxonomic rearrangements. In addition, we
provide revised diagnoses to the relevant genera based on
(1) osteological characters, (2) refining of species content,
and (3) comparison with another recently described
Southeast Asian arboreal toad (see Chandramouli et al.
2016) that was not available to Chan et al. (2016).
MATERIALS AND METHODS
Specimens (including types) of the taxa Pedostibes
tuberculosus, P. kempi,Rentapia hosii, R. everetti,R. rugosa,
and Bufoides meghalayanus were examined in collections
(see Appendix). Museum acronyms follow Sabaj Pe
(2014). We obtained comparable morphometric data and
distribution records from examined specimens, as well as
available literature (Dinesh and Radhakrishnan 2008; Frost
2015). With a dial caliper (60.1 mm) and a Leica Wild M3Z
dissecting microscope, we measured the following characters
(on the left side of the body for symmetric characters):
snout–vent length (SVL, from the tip of the snout to the
anterior margin of the cloaca), axilla–groin length (from the
posterior margin of the forelimb at its insertion point on
the body to the anterior margin of the hind limb at its
insertion point on the body), head length (HL, from the
posterior edge of the mandible to the tip of the snout), head
width (HW, the maximum width of the head at the angle of
the jaws), head depth (HD, the maximum depth of the head
at the region between the eye and the parotoid gland), body
width (the maximum width of the body at the trunk), eye
diameter (ED, the greatest horizontal diameter of the orbit),
eye–nostril length (EN, from the anterior border of the orbit
to the middle of the nostril), eye–snout length (ES, from the
anterior border of the orbit to the tip of the snout), snout–
nostril length (NS, from the tip of the snout to the middle of
the nostril), tympanum–eye length (TYE, from the posterior
border of the orbit to the anterior border of the tympanum),
upper eyelid width (UEW, the maximum width of the upper
eyelid), interorbital distance (IO, the shortest distance
between the dorso-medial margins of the orbits), internarial
distance (IN, the shortest distance between the dorsal
margins of the nostrils), tympanum diameter (TYD, the
greatest horizontal diameter of the tympanum), upper arm
length (on the dorsal surface, from the axilla to the inflection
of the flexed elbow), lower arm length (LAL, on the dorsal
surface, from the posterior margin of the elbow while flexed
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Herpetologica, 72(2), 2016, 137–147
E2016 by The Herpetologists’ League, Inc.
to the base of the outer metacarpal tubercle), palm length
(PAL, from the posterior border of the outer metacarpal
tubercle to tip of the longest finger), femur length (FEL,
from the anterior margin of the hind limb at its insertion
point on the body to the knee while flexed), tibia length
(TBL, from the posterior surface of the knee while flexed to
the base of the heel), foot length (FOL, from the base of the
inner metatarsal tubercle to the tip of the longest toe),
parotoid gland length (PL, the maximum length of the
parotoid gland), parotoid gland width (PW, the maximum
width of the parotoid gland), and finger and toe lengths
(from the tip of the disc to the nearest fork).
A subset of the above measurements (SVL, HL, HW, HD,
ED, EN, ES, IO, IN, TYD) was log transformed and
subjected to principal components analysis (PCA). Juvenile
specimens were excluded from the analysis. Statistically
informative tests were not performed on the holotype
specimen of Rentapia everetti because it is a subadult
specimen. However, extensive comparisons of its morpho-
logical characters were made. The data from the R.rugosa
holotype were used. The resulting component scores were
plotted to examine the distribution of the specimens in
multivariate morphological space. A distribution map of
these taxa was prepared based on relevant published
literature, type localities, and additional collection localities
of specimens examined and those of the corresponding
species listed in the VertNet database (available at http://
One specimen each of Pedostibes tuberculosus and
Bufoides meghalayanus was cleared and stained for studying
osteological features (following Hanken and Wassersug
1981). The clear-stained specimens were then examined
under an illuminated microscope in order to describe the
The PCA of the morphometric data extracted factors that,
taken together, explained 97.55% of the variance observed
between the samples and separated them into three discrete
clusters (Table 1; Fig. 1), each pertaining to a particular
genus from separate geographic regions. The first cluster
comprises the taxa Bufoides Pillai and Yazdani 1973 and
Pedostibes kempi from northeastern India. The second
cluster consists of only the type species, P. tuberculosus,
from the Western Ghats. The third cluster includes the
specimens of Rentapia hosii from different parts of Southeast
Asia (see Appendix) and the holotype of R. rugosa. Our
analysis of morphological data (Table 2) reveals that the
specimens allocated to the taxon P. tuberculosus show
a distinct morphological separation from P.kempi which,
in turn, clusters together with a morphologically similar and
geographically proximate taxon, Bufoides meghalayanus
(Yazdani and Chanda 1971) from Khasi Hills in the eastern
Himalayas. Rentapia hosii and R. rugosa collectively show
a morphological separation from the above two clusters
(P. tuberculosus and Bufoides +P. kempi).
In addition to the morphometric differences detailed
above, Pedostibes tuberculosus is distinguished from
P. kempi in having an exposed tympanum (vs. concealed in
P. kempi and Bufoides) and from both Rentapia hosii and
R. rugosa by the absence of a sharp tarsal fold (vs. present
and well-defined). Moreover, we observed the following
differences in osteological characters (Fig. 2). Pedostibes
tuberculosus and R. hosii have eight presacral vertebrae
while Bufoides meghalayanus has seven; sternum is bony in
P. tuberculosus while cartilaginous in R. hosii and B.
meghalayanus. Terminal phalanges in both fingers and toes
are widely expanded to truncate discs in P. tuberculosus,
relatively narrower in R. hosii, and with rudimentary
expansions in B. meghalayanus. Frontoparietal elements of
the skull are long and narrow in P. tuberculosus, trapezoidal
with a much broader posterior than anterior end in Rentapia,
and ovoid in Bufoides.
Based on our results and the above criteria, we hereby
restrict the genus Pedostibes to its type species, P.
tuberculosus, from the Western Ghats, southern India. Our
results show that the eastern Himalayan taxon, P. kempi,
exhibits morphological characters similar to, and diagnostic
of, the genus Bufoides Pillai and Yazdani 1973. Based on our
examination of the type specimens, we transfer P. kempi to
Bufoides. An examination of the types of the geographically
sympatric taxa Rentapia everetti and R. rugosua did not
reveal any morphological characters that serve to separate
these two nominal species. Therefore, we synonymize
Rentapia rugosa with R. everetti in accordance with the
principle of priority (ICZN 1999: Article 23).
Bufonidae Gray 1825
Pedostibes Gu¨ nther 1876
(Tables 2, 3; Figs. 2–4)
Type species.—Pedostibes tuberculosus Gu¨ nther 1876
Diagnosis (redefined herein).—The genus Pedostibes is
diagnosed by having small to moderate adult body size of
SVL 35.0–47.2 mm; absence of cephalic ridges; presence of
short, rounded parotoid glands; presence of an externally
visible tympanum; presence of eight presacral vertebrae;
a bony sternum; digit tips widely expanded to spatulate discs
with truncate anterior ends; short and broadly expanded
sacral diapophysis; absence of tarsal folds; glandular texture
of the dorsal skin; presence of partial webbing on fingers and
complete webbing on toes (Table 2).
Comparison.—The genus, Pedostibes, a member of the
largely South Asian Adenominae clade (Bocxlaer et al. 2009),
is distinguished from Rentapia and other Asian bufonid
genera (comparative traits in parentheses) by having smaller
adult body size of SVL 35.0–47.2 mm (52.3–99.5 mm in
Rentapia; up to 215 mm in Phrynoidis [fide Malkmus et al.
2002]); absence of cephalic ridges (present in some species
of Duttaphrynus,Adenomus, and Xanthophryne); presence
of short, rounded parotoid glands (absent in Sabahphrynus);
presence of an externally visible tympanum (concealed in
Sabahphrynus and Bufoides); presence of partial webbing on
fingers (absent in Duttaphrynus,Xanthophryne,Adenomus,
Ghatophryne, and Phrynoidis) and complete webbing on
toes (incomplete webbing in Sabahphrynus); absence of
a tarsal ridge (present in Rentapia); presence of eight
presacral vertebrae (vs. seven in Bufoides); and glandular
texture of the dorsal skin (smooth or granular in Rentapia,
Xanthophryne,Adenomus, Ghatophryne,Bufoides, and
138 Herpetologica 72(2), 2016
Redescription based on a voucher specimen (Wildlife
Institute of India, Dehradun, India [WII] 38.6.91 from
Tamil Nadu, India).—An unsexed adult, SVL 47.2 mm.
Head slightly convex, smooth, and wider than long (HL
84.0% of HW, 25.0% of SVL), lacking cranial ridges. Nostril
oval in shape, laterally oriented, with a slender inner margin.
Internarial area smooth and concave. Canthal ridges weakly
defined and rounded in dorsal aspect. Snout smooth, flat in
lateral aspect and pointed in dorsal aspect (ES 35.8% of HL,
9.0% of SVL). Nostril to snout distance less than distance
between eye and nostril (NS 3.0% of EN). Loreal region
oblique and smooth. Large distinctly visible tympanum,
vertically oval. Parotoid glands weakly distinct, rounded,
small, smooth and undivided (unlobulated). Interorbital area
flat, broad and smooth, IN less than interorbital width (IN
52.8% of IO). Upper eyelids concave, their outer edges
rounded, width less than interorbital width.
Lower arm shorter than head (LAL 112.0% of HL, 28.0%
of SVL). Fingers with basal webbing and tips of fingers sharp
and smooth, Finger III the longest. Finger I slightly shorter
than II. Relative length of fingers I ,II ,IV ,III.
Terminals with widely expanded discs; base of the Finger IV
with a large, flat, oval outer metacarpal tubercle.
Femur and tibia equal in length (FEL 46.0% of SVL).
Foot length shorter than TBL (FOL 104.5% of TBL, 47.0%
of SVL). Two metatarsal tubercles present, a larger lateral
tubercle and a smaller medial tubercle; tarsal ridge absent.
Toe IV the longest. Toes completely webbed; the dorsal and
ventral surfaces of all toes smooth, their tips rounded.
Skin structures.—Dorsal skin glandular in texture; not
warty. Small globular glandules present on the dorsal and
dorso-lateral regions, more intensely on posterior than on
anterior part. Ventral surface with small warts, less dense on
anterior and posterior ends but intense on the belly. Groin
and undersurfaces of the thigh smooth.
Coloration (in preservative).—Pale brown above with
dark and white marbled pattern along the flanks; ventral
surfaces with marbled pattern.
Distribution.—Pedostibes is endemic to the Western
Ghats of Peninsular India and has been recorded from
localities between the Agasthyamalai Hills (8.50uN, 77.47uE,
1000 m elevation; in all cases, datum 5WGS84) in the
south, northward to Amboli (15.96uN, 73.99uE, 744 m
elevation) in the northern portion of the Western Ghats
(Fig. 4; Dinesh and Radhakrishnan 2008).
Included species.—Pedostibes tuberculosus Gu¨ nther
1876 (with the genus Pedostibes being monotypic).
Rentapia Chan, Grismer, Zacharia, Brown, and Abraham
(Tables 2, 3; Figs. 2–4)
Type species.—Nectophryne hosii Boulenger 1892.
Diagnosis.—Rentapia is characterized by having large
adult body size of SVL 52.3–99.5 mm, long and relatively
narrow sacral diapophyses, and presence of a cartilaginous
sternum (Table 2). In addition to the traits described by
Chan et al. (2016), this genus is characterized by the
presence of tarsal folds and smooth or granular texture of the
Comparison.—Phylogenetically, the genus Rentapia is
not a member of the Adenominae clade and can be defined
as the member of a clade including Phrynoidis as its closest
taxon (Matsui et al. 2007, 2015; Bocxlaer et al. 2009; Pyron
and Wiens 2011; Ron et al. 2015; Chan et al. 2016), but not
Pedostibes tuberculosus.Rentapia is distinguished from
Pedostibes,Phrynoidis, and other Asian bufonid genera
(characters in parentheses) by having larger adult body size
of SVL 52.3–99.5 mm (21.8–27.4 mm in Blythophryne
Chandramouli, Vasudevan, Harikriahnan et al. 2016; 35.0–
47.2 mm in Pedostibes; 215 mm in Phrynoidis [fide Malkmus
et al. 2002]); absence of cephalic ridges (present in some
species of Duttaphrynus,Adenomus,andXanthophryne);
presence of short, rounded or triangular parotoid glands
(absent in Sabahphrynus, slender and elongated in Blytho-
phryne); presence of an externally visible tympanum (hidden
in Sabahphrynus and Bufoides); presence of expanded discs
in finger and toe tips (absent in Phrynoidis); presence of
TABLE 1.—Factor loadings and the percentage of variance explained by
a principal components analysis of morphometric values from five species of
arboreal toads in Southeast Asia.
SVL 0.33 0.00 20.01 0.02 0.09
HL 0.33 20.02 20.20 20.41 20.06
HW 0.33 0.06 20.19 20.43 0.08
HD 0.32 20.29 20.25 0.13 20.33
ED 0.32 20.19 20.39 20.02 0.38
EN 0.31 0.12 0.58 20.06 20.44
ES 0.33 0.10 0.21 20.32 20.14
IO 0.31 20.12 0.52 0.18 0.67
IN 0.29 0.82 20.23 0.42 0.02
TYD 0.31 20.41 20.04 0.56 20.26
Eigenvalue 8.88 0.35 0.22 0.17 0.15
% variance 88.77 3.47 2.18 1.66 1.47
SVL 5snout–vent length, HL 5head length, HW 5head width, HD 5head depth, ED 5eye
diameter, EN 5eye–nostril length, ES 5eye–snout length, IO 5interorbital distance, IN 5
internarial distance, TYD 5tympanum diameter.
TABLE 2.—Comparison of the characters of the morphologically similar species of arboreal toads formerly assigned to the genus Pedostibes.
Character Pedostibes tuberculosus (n53) Rentapia hosii (n543) Rentapia everetti (n52) Bufoides kempi (n52)
Type locality “Malabar,” South India Mt. Dulit, Borneo Mt. Kinabalu, Borneo Tura, Garo Hills, Northeast India
Adult snout–vent length (mm) 35.0–47.2 52.3–99.5 74.3 (n51) 17.3–29.8
Cephalic ridges Absent Absent Absent Present
Parotoid glands Short and round Elongated Oval Elliptical
Tympanum Distinct Distinct Moderately distinct Hidden
Tarsal folds Absent Present Present Absent
Dorsal skin Glandular Smooth Granular Granular
Webbing on fingers Partial Basal Basal Basal
Webbing on toes Fully Fully Fully 3/4
Throat Granular Granular Smooth Granular
CHANDRAMOULI AND AMARASINGHE—TAXONOMY OF PEDOSTIBES 139
partial webbing on fingers (absent in Phrynoidis) and complete
webbing on toes (incomplete webbing in Sabahphrynus);
presence of eight presacral vertebrae (vs. seven in Bufoides
and six in Blythophryne); presence of a tarsal ridge (absent in
Pedostibes and Bufoides); and smooth or rugose texture of the
dorsal skin (granular in Phrynoidis; glandular in Pedostibes).
Distribution.—Rentapia occurs in Peninsular Thailand,
Malaysia, Borneo, and Sumatra (Fig. 4; Frost 2015; Chan
et al. 2016).
Included species.—Rentapia hosii (Boulenger 1892) and
R. everetti (Boulenger 1896).
Rentapia hosii (Boulenger 1892)
Nectophryne hosii Boulenger 1892:508.
Pedostibes hosii––Barbour (1938:192).
Pedostibes hosi––Inger (1958:478).
Pedostibes hosei––Inger (1966:93).
Rentapia hosii—Chan et al. 2016:9.
Holotype of Nectophryne hosii.—Adult male (Natural
History Museum, London, UK [BMNH] 1922.214.171.124;
formerly 126.96.36.199), “Mt. Dulit, Sarawak,” East Malaysia
Diagnosis.—Rentapia hosii is distinguished by having
large adult body size of SVL 52.3–99.5 mm, absence of
cephalic ridges, presence of elongated parotoid glands,
presence of an externally visible tympanum, presence of
tarsal folds, smooth texture of the dorsal skin, presence of
basal webbing on fingers and complete webbing on toes, and
granular texture of the throat skin (Table 2).
FIG. 1.—Plotted results of principal component (PC) analyses showing morphometric separation between five species of arboreal toads: Pedostibes
tuberculosus (open diamonds), Rentapia hosii (filled circles), R. everetti (open circles), Bufoides meghalayanus (open squares), and B. kempi (filled squares).
(A) PC1 vs. PC2; (B) PC1 vs. PC3; (C) PC1 vs. PC4; (D) PC1 vs. PC5.
140 Herpetologica 72(2), 2016
FIG. 2.—Cleared and stained specimens of Pedostibes tuberculosis (first column), Rentapia hosii (second column), and Bufoides meghalayanus (third
column) showing dorsal views of the following skeletal characters: skull and vertebral column (A–C), pectoral girdle (D–F), feet (G–I), and hand (J–L). FP 5
frontoparietal; SD 5sacral diapophysis; CL 5clavicle; COR 5coracoid; ST 5sternum. A color version of this figure is available online.
CHANDRAMOULI AND AMARASINGHE—TAXONOMY OF PEDOSTIBES 141
Redescription of holotype.—An adult male SVL of 59.8
mm. Head slightly concave, smooth and wider than long (HL
96.7% of HW, 29.9 of SVL). Nostril oval in shape, laterally
oriented, with a slender inner margin. Internarial area
smooth and concave. Canthal ridges prominently defined
and rounded in dorsal aspect. Snout smooth, rounded in
lateral aspect and pointed in dorsal aspect (ES 39.1% of HL,
11.7% of SVL), NS less than distance between eye and
nostril (NS 34.6% of EN). Loreal region oblique and smooth.
Large distinctly visible tympanum, vertically oval (TYE 4.5%
of HL). Parotoid glands elongated, narrow (posteriorly
wider), smooth, and undivided (unlobulated; PW 46.4% of
PL; PL 38.5% of HL). Interorbital area concave and smooth,
IN less than interorbital width (IN 44.8% of IO). Upper
eyelids concave, their outer edges rounded, width less than
interorbital width (UEW 58.9% of IO).
Lower arm slightly shorter than head (LAL 96.6% of HL,
28.9% of SVL). Fingers with basal webbing and tips of
fingers sharp and smooth, Finger III the longest (FL3 66.8%
of PAL, 59.5% of LAL, 147.1% of ES). Finger I slightly
shorter than II (FL1 72.7% of FL2). Relative length of
fingers I ,II ,IV ,III, a nuptial pad on dorsal side of
Finger I. Subarticular tubercles on fingers rounded, two
palmar tubercles, the lateral one larger than the medial.
Femur and tibia equal in length (FEL 48.1% of SVL).
Foot length shorter than TBL (FOL 88.2% of TBL, 42.5% of
SVL). Two metatarsal tubercles present, the medial one oval,
the lateral one rounded; tarsal ridge present. Toe IV the
longest (TL4 55.5% of FOL, 78.7% of HL, 201.4% of ES,
167.8% of TL3). Toes completely webbed, the dorsal and
ventral surfaces of all toes smooth, their tips rounded.
Relative length of toes I ,II ,III ,V,IV.
Skin structures.—Throat rough and granulated, but belly
smooth. Both sides of the gape without warts, and distal
edges of lower and upper jaws are smooth. Anterior end of
mandible with symphysial knob. Axila-to-groin area smooth.
Middorsal area smooth and larger warts present on lateral
and posterior parts of the body. Both dorsal and ventral
surfaces of arms covered with sharp and rough small warts
on dorsal aspect. Thigh relatively smooth on both anterior
and posterior surfaces.
Coloration (in preservative).—Dorsum uniform with
olive brown; dorsal surfaces of limbs without any bars; all
ventral surfaces similar to dorsal coloration, except for
a darker throat.
Variation.—In some live specimens from East Kaliman-
tan, we observed a dark variegated skin color pattern in
Distribution.—Rentapia hosii occurs in peninsular Thai-
land, Malaysia, Borneo, and Sumatra (Fig. 4; Frost 2015).
Rentapia everetti (Boulenger 1896)
Nectophryne everetti Boulenger 1896:450.
Pedostibes everetti—(Barbour 1938:192).
Pedostibes rugosus Inger 1958:476–478 syn. nov.
Rentapia rugosus—Chan et al. 2016:9,11.
Rentapia rugosa—Chandramouli and Amarasinghe (2016)
Holotype of Nectophryne everetti.—A subadult male
(BMNH 19188.8.131.52; formerly 184.108.40.206), “Mount Kina
Balu, North Borneo,” Sabah, East Malaysia (Borneo).
Nomenclatural note.—When transferring the taxon
Pedostibes rugosus Inger 1958 to the genus Rentapia, Chan
et al. (2016) used the combination Rentapia rugosus (sic.).
According to the International Code of Zoological Nomen-
clature (ICZN 1999) Articles 30 and 31, the generic name
Rentapia erected by these authors is feminine in gender.
Because the specific epithet, rugosus, is an adjective in
masculine gender, referring to the rugose texture of its skin,
we amend it here as Rentapia rugosa, in accordance with the
Article 31.2 (Agreement in Gender).
Diagnosis.—Rentapia everetti is distinguished by having
large adult body size of SVL 74.3 mm, absence of cephalic
ridges, presence of oval parotoid glands, presence of an
externally visible tympanum, presence of tarsal folds,
granular texture of the dorsal skin, presence of basal
webbing on fingers and complete webbing on toes, and
smooth texture of the throat skin (Table 2).
Redescription of the holotype of Nectophryne ever-
etti.—A subadult male SVL of 33.0 mm. Head slightly
concave, wider than long (HW 118.5% of HL, 34.8% of SVL)
and with numerous tiny round warts. Nostril round in shape,
laterally oriented, with a slender inner margin, Internarial
area concave with tiny warts. Canthal ridges prominently
defined and sharp in dorsal aspect. Snout with tiny warts, flat
in both lateral and dorsal aspects (ES 34.8% of HL, 12.1% of
SVL), Nostril to snout distance less than distance between
eye and nostril (NS 44.0% of EN). Loreal region oblique and
TABLE 3.—Comparison of the morphometric measurements (mm) of
the holotypes of Rentapia hosii,R. everetti and its junior synonym,
Rentapia hosii Rentapia everetti
Snout–vent length 59.8 33.0 74.3
Axilla–groin length 29.8 14.7 24.5
Head length 17.9 11.5 22.7
Head width 18.5 9.7 24.8
Head depth 10.3 5.6 11.0
Body width 13.5 7.9 17.5
Eye diameter 6.9 4.3 6.5
Eye–nostril length 5.2 2.5 6.3
Snout–nostril length 1.8 1.1 2.5
Eye–snout length 7.0 4.0 9.7
Tympanum–eye length 0.8 0.8 1.1
Upper eyelid width 4.6 2.7 8.5
Interorbital distance 7.8 3.7 7.6
Internarial distance 3.5 1.5 4.5
Tympanum diameter 3.5 1.3 3.2
Upper arm length 11.3 7.0 19.8
Lower arm length 17.3 8.2 21.4
Palm length 15.4 10.6 24.6
Femur length 28.8 11.5 30.4
Tibia length 28.8 15.4 33.5
Foot length 25.4 12.9 34.2
Parotoid gland length 6.9 4.3 10.1
Parotoid gland width 3.2 1.7 7.4
Finger I length 4.8 1.5 5.0
Finger II length 6.6 3.7 11.9
Finger III length 10.3 5.5 16.2
Finger IV length 9.5 2.6 10.7
Toe I length 3.8 2.1 5.7
Toe II length 5.5 2.5 8.6
Toe III length 8.4 3.7 11.4
Toe IV length 14.1 6.1 18.9
Toe V length 9.5 4.3 11.7
142 Herpetologica 72(2), 2016
smooth. Small distinctly visible tympanum, vertically oval
(TYE 6.9% of HL). Parotoid glands elongated, narrow
(posteriorly wider), rough and undivided (unlobulated; PW
39.5% of PL; PL 37.4% of HL). Interorbital area flat and
with tiny warts, internarial distance less than interorbital
width (IN 40.5% of IO). Upper eyelids concave, their outer
edges sharp, width less than interorbital width (UEW 72.9%
Lower arm shorter than head (LAL 71.3% of HL, 24.8%
of SVL). Fingers with basal webbing, web reaching
subarticular tubercle of Finger I, beyond basal subarticular
tubercle on lateral edge of II. Tips of fingers sharp and
smooth, with Finger III being the longest (FL3 51.8% of
PAL, 67.1% of LAL, 137.5% of ES). Finger I shorter than II
(FL1 40.5% of FL2). Relative length of fingers I ,II ,IV
,III, a nuptial pad on dorsal side of Finger I. Subarticular
tubercles on fingers rounded; two palmar tubercles, the
lateral one larger than the medial.
Femur shorter than tibia in length (FEL 34.8% of SVL).
Foot length shorter than TBL (FOL 83.7% of TBL, 39.1% of
SVL). Two metatarsal tubercles present, the medial one oval,
the lateral one rounded; tarsal ridge present. Toe IV the
longest (TL4 47.3% of FOL, 53.0% of HL, 174.3% of ES,
164.8% of TL3). Toes completely webbed, the dorsal and
ventral surfaces of all toes smooth, their tips rounded.
Relative length of Toes I ,II ,III ,V,IV.
Skin structures.—Throat smooth and not granulated,
belly smooth. Both sides of the gape without warts, and distal
edges of lower and upper jaws are smooth. Anterior end of
mandible with symphysial knob. Axila-to-groin area is
smooth. Middorsal area, lateral and posterior parts of the
body present with larger warts. Both dorsal and ventral
surfaces of arms covered with sharp and rough small warts
on dorsal aspect. Thigh comparatively smooth on both
anterior and posterior surfaces.
Coloration (in preservative).—Dorsum bluish grey and
limbs light brown; back with many dark brown blotches,
lateral surfaces of head with broad, dark brown bars; dorsal
surfaces of limbs cross barred; all ventral surfaces immac-
Distribution.—Rentapia everetti occurs in Borneo
(Fig. 4; Frost 2015).
FIG. 3.—Dorsal aspect of the bodies of five species of arboreal toads collected from southern and Southeast Asia: (A) holotype of Rentapia hosii BMNH
19220.127.116.11; (B) holotype of R. everetti BMNH 1918.104.22.168; (C) holotype of Rentapia rugosa (junior synonym of R. everetti) FMNH 81297; (D) a syntype
of Pedostibes tuberculosus BMNH 1922.214.171.124; (E) a voucher specimen of Bufoides meghalayanus (WII uncatalogued); and (F) a syntype of Nectophryne
kempi (ZSI 18481A). In each panel, bar 55 mm. A color version of this figure is available online.
CHANDRAMOULI AND AMARASINGHE—TAXONOMY OF PEDOSTIBES 143
Bufoides Pillai and Yazdani 1973
(Table 2; Figs. 2–4)
Type species.—Ansonia meghalayana Yazdani and
Diagnosis (redefined herein).—A genus of small-
bodied (SVL 29.8–47.2 mm), rupicolous toads, characterized
by presence of supraorbital, preorbital, and postorbital
ridges on the top of the head; presence of short, oval-
shaped parotoid glands; absence of an externally visible
tympanum; digit tips with rounded, poorly dilated discs;
presence of seven presacral vertebrae; presence of a carti-
laginous sternum; granular texture of the dorsal skin;
absence of tarsal folds; and presence of basal webbing on
fingers and complete webbing on toes (Table 2).
Comparison.—Bufoides can readily be distinguished
from other oriental bufonid genera (characters in parenthe-
ses) by the absence of an externally visible tympanum
(exposed in Pedostibes,Blythophryne,Parapelophryne, and
Rentapia); presence of parotoid glands (absent in Parapelo-
phryne and Pelophryne); presence of cranial ridges (absent
in Pedostibes,Blythophryne,Parapelophryne, and Renta-
pia); poorly developed digital discs on fingers and toe tips
(well developed, widely dilated discs in Pedostibes,Blytho-
phryne, and Rentapia); presence of seven presacral verte-
brae (six in Blythophryne, eight in Pedostibes,Parapelo-
phryne,andRentapia); granular skin texture (glandular in
Pedostibes); and the absence of tarsal folds (present in
Distribution.—Bufoides in restricted in distribution to
the Garo and Khasi hills of the eastern Himalayas, with one
species occurring on each of these hill ranges.
Included species.—Bufoides kempi (Boulenger 1919)
comb. nov. and Bufoides meghalayanus (Yazdani and
Bufoides kempi (Boulenger 1919) comb. nov.
(Table 2; Figs. 3, 4)
Nectophryne kempi (Boulenger 1919:207).
Pedostibes kempi—Barbour (1934:192); Chan et al. (2016:13).
FIG. 4.—Collection localities for five species of arboreal toads in Southeast Asia, modified after Chan et al. (2016): Bufoides kempi (open square, type
locality), B. meghalayanus (closed square, type locality), Pedostibes tuberculosus (closed circles), Rentapia everetti (open triangle, type locality of R. rugosa
with a dot in the middle), and R. hosii (open circles, type locality with a dot in the middle).
144 Herpetologica 72(2), 2016
Diagnosis.—Bufoides kempi is diagnosed by small to
moderate body size (SVL 17.4–29.8 mm); presence of
nonkeratinized cranial ridges; presence of short, oval-shaped
parotoid glands; absence of an externally visible tympanum;
moderate degree of webbing on toes and basal webbing on
fingers, with poorly developed terminal discs without lateral
Redescription of the syntypes.—Characters of Zoolog-
ical Survey of India, Kolkata, India (ZSI) 18481A followed,
when appropriate, by those of ZSI 18481B in parentheses.
An adult and a subadult (unsexed) specimen, each measuring
29.8 mm (17.4 mm) SVL; head distinct from body, broader
than long, HL 89.0% (87.0%) of HW, 30.0% (33.0%) of SVL.
Nostril to snout distance less than distance between eye and
nostril, NS 33.0% (23.0%) of EN. Snout rounded in dorsal
and truncate in lateral views; canthal fold not pronounced.
Loreal region flat; top of the snout concave; tympanum
absent; a large conical tubercle present at the jaw angle.
Parotoid glands lobulated, short, and bean-shaped, extend-
ing from the postorbital region to the axillary region on the
dorsum. Interorbital space broader than internarial distance,
IN 49.0% (64.0%) of IO. Upper eyelids with dense granules;
narrower than interorbital distance.
Lower arm longer than head, LAL 139.0% (115.0%) of
HL, 24.0% (26.0%) of SVL. Fingers webbed at the base,
fingertips with small discs at tips, damaged and broken in the
larger specimen. A large outer palmar tubercle present at the
base of the palm.
Femur longer than tibia in the smaller specimen, FEL
37.0% (43.0%) of SVL, tibia broken in the larger syntype.
Two oval metatarsal tubercles present; inner one larger than
outer; tarsal ridge not discernible. Most of the toes damaged
in both the specimens but the left foot of the smaller
specimen shows partial webbing between Toes II and III,
extending half the length of Toe III and similarly on Toe IV;
toe tips with small discs.
Skin structures.—Dorsum granular with small conical
warts scattered throughout; more intense toward lateral area
than on the dorsal region. Venter with similar granules of
smaller size; more intense on the posterior than on anterior end.
Coloration in preservative.—The larger of the two
syntypes olive brown above with dark black flanks; the
smaller specimen uniform black throughout; undersides
without pattern, black in both specimens.
Distribution.—This species is known only from its type
locality, Tura, in the Garo Hills of northeastern India.
Bufoides meghalayanus (Yazdani and Chanda 1971)
Ansonia meghalayana Yazdani and Chanda 1971.
Diagnosis.—Bufoides meghalayanus is diagnosed by
small to moderate adult size (SVL 31.2–47.2 mm); presence
of pre-, post-, and supraorbital ridges on the head; presence
of short, oval-shaped parotoid glands; absence of an
externally visible tympanum; digit tips with rounded, poorly
dilated discs; granular texture of the dorsal skin; absence of
tarsal folds; presence of basal webbing on fingers and nearly
complete webbing on toes, with poorly developed terminal
Description of a voucher specimen (WII uncatalo-
gued, from Khasi Hills, Meghalaya, India).—An
unsexed adult, SVL of 31.2 mm. Head depressed, a little
broader than long (HL 110.0% of HW, 35.0% of SVL);
snout projecting beyond mandible, with a rounded tip in
both dorsal and lateral views (ES 39.0% of HL, 14.0% of
SVL); nostrils pointing downward, located midway be-
tween eyes, a little closer to snout tip than to the eyes (NS
202.0% of EN); loreal region slightly concave with an
indistinct canthal fold. Preorbital, postorbital, and supra-
orbital ridges on the head prominent; not keratinized.
Loreal and internarial regions concave. Tympanum absent;
parotoid glands slender and slightly elongate (PW 13.0%
of PL; PL 48.0% of HL); extending downward from the
postorbital ridge. Interorbital distance greater than upper
eyelid width (IN 58.0% of IO).
Lower arm much shorter than the head (LAL 70.0% of
HL); palm as long as the upper arm, with fleshy webbing
extending only to the base of the fingers; relative lengths of
fingers I ,II ,IV ,III. Subarticular tubercles on fingers
indistinct, outer palmar tubercle large and distinct.
Femur a little longer than the tibia (FEL 43.0% of SVL);
foot nearly as long as tibia length (FOL 97.0% of TBL, 39%
of SVL). Two metatarsal tubercles present, the medial one
oval, the lateral one rounded; tarsal ridge absent. Toes with
well-developed webbing; their relative lengths I ,II ,III
,V,IV; digit tips with small fleshy discs without expanded
Skin structures.—Dorsal surfaces with small, almost
uniform granular warts scattered all over the body; arms and
legs. Ventral surfaces rough with granules sparsely distrib-
uted in the anterior region from the chin, across the throat to
the axilla; densely granulated posteriorly.
Coloration in preservative.—Dorsum without pattern,
uniform greyish brown; venter pale white in color throughout.
Distribution.—Bufoides meghalayanus has been recorded
from a few localities in the Khasi Hills of Meghalaya near
Mawblang, Cherrapunji. Other records from this region were
provided by Deuti et al. (2012).
After Boulenger (1892, 1896) described Nectophryne hosii
and N. everetti, Inger (1958) described Pedostibes rugosus
from “Menuang, headwaters of the Baleh River” in Sarawak,
based on a male holotype (Field Museum of Natural History,
Chicago, USA [FMNH] 81297) and a female paratype. This
taxon was diagnosed solely on the presence of “large oval
parotoid glands, numerous round warts dorsally, and a sharp
tarsal fold.” Upon describing this taxon, Inger (1958)
remarked that only two species of Pedostibes,P. hosii and
P. rugosus, have a sharply defined tarsal fold; thus, he
eliminated P. tuberculosus from further comparisons and
distinguished P. rugosus from P. hosii based on a set of
characters. Two more congeners (after Barbour 1938), P.
kempi and P. everetti, were neither mentioned nor
compared. Among them, P. kempi is easily distinguishable
from the rest of the species by lacking an externally visible
tympanum (Boulenger 1919).
The diagnostic characters of Pedostibes rugosus in its
original description coincide with those described for the
taxon P. everetti by Boulenger (1896), which was apparently
overlooked. Additionally, the type localities of these two taxa
(i.e., “Menuang, headwaters of the Baleh River” for P.
rugosus and “Mt. Kina Balu, north Borneo” for P. everetti)
CHANDRAMOULI AND AMARASINGHE—TAXONOMY OF PEDOSTIBES 145
are situated within the same biogeographical region of
Borneo (Fig. 4). Malkmus et al. (2002) remarked on the
similarity between P. rugosus and P. everetti, and cited
Manthey and Grossman (1997), who suggested that none of
the diagnostic characters of P. rugosus were distinct from
character states exhibited by P. everetti.AlthoughInger
(1966) mentioned additional morphological differences be-
tween P. rugosus and P. everetti, our reexamination of their
type specimens leads us to conclude that Inger’s (1966)
characters are not sufficiently diagnostic to distinguish those
species. This is further supported by the fact that P. everetti
has seldom been mentioned in field-based scientific studies
since the name Pedostibes rugosus came into being, essentially
concealing the former taxon (e.g., see Das 2007; Matsui et al.
2007, 2015; Pyron and Wiens 2011; Ron et al. 2015). Our
reexamination of the type specimens (BMNH 19126.96.36.199
and FMNH 81297, respectively) and original descriptions
of these two taxa support Manthey and Grossman (1997) in
that the morphological distinction between P.rugosus and
P.everetti is effectively absent.
In the original description of Nectophryne kempi,Boulen-
ger (1919) distinguished it from “congeners” based on the
following diagnostic characters: tympanum hidden; fingers
with feebly dilated, truncated tips, 1/3 webbed; toes 3/4
webbed, tips rounded but not dilated; two small metatarsal
tubercles, absence of tarsal fold; presence of prominent
parotoid glands; and large, yellowish axillary spots. He also
remarked on the similarity of N. kempi with N. maculata in
lacking an externally visible tympanum (Boulenger 1919).
Barbour (1938: 192) transferred N. kempi to Pedostibes with
some degree of caution by stating “The four species of
Pedostibes first mentioned and possibly this fifth one as well
[P. kempi] are alike…” where he refers to the taxa P.
tuberculosus,P.hosii,P.everetti,andP. altitudinis [the
lattermost of which currently represents a synonym of an
unrelated species, Ansonia fuliginea (Mocquard 1890); see
Frost 2015]. Additionally, even after revising Pedostibes,Chan
et al. (2016) attributed the “Indian” species kempi to this
genus, making a remark on the uncertainty of this allocation.
Moreover, limited taxon sampling has hindered Chan et al.
(2016) from comparing this taxon with the other arboreal toad
genus Bufoides from the eastern Himalayan region.
When erecting it for the taxon Ansonia meghalayana
Yazdani and Chanda 1971, Pillai and Yazdani (1973)
characterized the genus Bufoides based on the presence of
cranial ridges, unwebbed fingers with dilated tips, almost
fully webbed toes, and a concealed tympanum. Das et al.
(2009) procured further specimens of B. meghalayanus from
the type locality and identified an unstudied diversification
within Bufoides endemic to India by referring to specimen
MFA 10134 collected from Tura, Garo Hills (the type
locality of Nectophryne kempi), which is not conspecific with
B. meghalayanus. Prompted by its provenance, we suspect
that this specimen could possibly represent the taxon kempi.
Furthermore, Das et al. (2009) identified a discrepancy from
the original description, that the type series of B. megha-
layanus has well-developed parotoid glands. Thus, both the
taxa Pedostibes kempi and B. meghalayanus share the
following suite of morphological characters considered to
be diagnostic: absence of an externally visible tympanum,
presence of parotoid glands, and well-developed webbing in
toes. The two species occur in two different hill ranges,
however: B. kempi from Tura, Garo Hills and B. megha-
layanus from Mawblang, Khasi Hills. Our reexamination the
specimens of these two taxa complies with the above-
mentioned set of similarities. Hence, we transferred the
taxon Pedostibes kempi to the genus Bufoides in the new
combination, Bufoides kempi. Whether B. meghalayanus and
B. kempi represent the same species or not, however, is
a question that remains open for further research. Pending
collection of new specimens referable to P. kempi from the
Garo Hills, we take a conservative stand and retain them to
be specifically distinct.
The systematic status of the South and Southeast Asian
bufonids at the generic level has long been problematic, as
interpreted from the following examples. Once considered to
be globally distributed, the genus Bufo Garsault 1764 was
subjected to several systematic and taxonomic studies
resulting in the reevaluation and recognition of several valid
genera such as Adenomus,Duttaphrynus,Ingerophrynus,
Phrynoidis,Vandijkophrynus, and Xanthophryne (see Man-
amendra-Arachchi and Pethiyagoda 1998; Frost et al. 2006;
Biju et al. 2009). All of these clades show a narrow, finite
geographic distribution range (Adenomus: Sri Lanka; Dut-
taphrynus,Ingerophrynus, and Phrynoidis: South and
Southeast Asia; Vandjikophrynus: South Africa; and Xantho-
phryne: northern portion of the Western Ghats in peninsular
India; Frost 2015). Similar cases are also known for certain
genera that were once considered to be widely distributed,
with some species showing disjunct distribution ranges.
Examples include the transfer of Ansonia ornata Gu¨ nther
1876 to Ghatophryne by Biju et al. (2009), and validation of
Pedostibes from the synonymy of Nectophryne Buchholz and
Peters 1875 by Barbour (1938). Frost (2015) suggested
a similar scenario for Pedostibes according to its current
definition by stating that, instead of providing a taxonomic
remedy, Pyron and Wiens (2011) embraced a polyphyletic
Pedostibes. Ron et al. (2015) also commented on the
problem of a polyphyletic Pedostibes, and suggested generic
transfer of the taxa hosii and rugosus to Phrynoidis, although
they were not transferred formally. Chan et al. (2016)
resolved this problem by erecting a new generic name
Rentapia for the Sundaic species hosii,everetti,andrugosus.
Because Chan et al. (2016) did not present a comprehensive
taxon-sampling (e.g., Bufoides) or examination of the name-
bearing types, they provided some tentative conclusions at
both generic and specific levels in some cases. Chan et al.
(2016) expressed uncertainty with such species and thus
made some provisional rearrangements within this clade.
Our reexamination of the types and other material has
revealed finer patterns within this group, and led to a few
other taxonomic rearrangements.
Acknowledgments.—SRC thanks Wildlife Institute of India for the award of
a junior research fellowship. AATA thanks the Ministry of Research and
Technology of the Republic of Indonesia, particularly S. Wahyono and L.
Shalahuddin, for granting research permits; R. Ubaidillah, A. Hamidy,
Syaripudin, W. Trilaksana, and other staff members of Museum Zoologicum
Bogoriense, Bogor, Indonesia (MZB), and P.D. Campbell of BMNH for
facilitating in-house study of specimens under their care. We also thank J.
Supriatna and the staff of the Research Center for Climate Change, University
of Indonesia, for their support; A. Resetar and R. Grill (FMNH), P.D.
Campbell and J. Streicher (BMNH), V. Thakur (vertebrate museum, WII),
K. Venkataraman (ZSI), K. Chandra (ZSI), and K. Deuti (ZSI) for access,
photographs, and data about the specimens under their care. Finally, we thank
the Editor, C.P. Groves, and anonymous reviewers for valuable comments.
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Accepted on 27 January 2016
Associate Editor: Christopher Raxworthy
Bufoides kempi (Boulenger 1919).—Garo Hills, Assam, Meghalaya, India:
ZSI 18481A–B (syntypes).
Bufoides meghalayanus (Yazdani and Chanda 1971).—Meghalaya,
Northeast India: WII uncatalogued.
Pedostibes tuberculosus Gu¨ nther 1876.—Malabar (Western Ghats), South
India: BMNH 19188.8.131.52–71 (syntypes); Tamil Nadu, South India: WII
Rentapia everetti (Boulenger 1896).—Mt. Kina Balu, Sabah, Malaysia:
BMNH 19184.108.40.206 (holotype); Menuang, headwaters of the Baleh River,
Third Division, Sarawak, Malaysia: FMNH 81297 (holotype of Pedostibes
rugosus Inger 1958).
Rentapia hosii (Boulenger 1892).—Mt. Dulit, Sarawak, Malaysia: BMNH
19220.127.116.11 (holotype); East Kalimantan, Indonesia: MZB 7779, 7780,
7864–67, 8891–99, 8900, 15454, 15455; Central Kalimantan, Indonesia:
3127, 3136, 8901–05, 10804, 10805, 11777, 11778, 17098; West Kalimantan,
Indonesia: 4758, 4759, 4761, 7250; South Kalimantan, Indonesia: 6083–89;
Aceh, Sumatra, Indonesia: 6904, FMNH 77369.
Phrynoidis juxtasper (Inger 1966).—N. Borneo, Tauwau Dist, Kalabakan,
Sungei Kadat: Museum of Comparative Zoology, Harvard, USA (MCZ) A-
CHANDRAMOULI AND AMARASINGHE—TAXONOMY OF PEDOSTIBES 147