RESEARCH ARTICLE
2017 | VOLUME 33 | ISSUE 1-4 | PAGES 47-54
Rediscovery and redescription of a little known,
insular endemic frog, Ingerana charlesdarwini
(Das, 1998) (Amphibia: Anura: Dicroglossidae) from
the Andaman Islands, Bay of Bengal
S. R. Chandramouli1*
1.
Wildlife Institute of India, Chandrabani, Dehradun, Uttarakhand, 248001, India
The critically endangered, insular endemic dicroglossid frog Ingerana charlesdarwini was described
from three adult specimens from Mount Harriet National Park, South Andaman (India). The description
of this species is here expanded based on fresh collections made from the Andaman archipelago in the
Bay of Bengal. Morphometric analysis based on a larger sample size of fourteen specimens including
the ones collected during this study revealed that this species is not sexually dimorphic or dichromatic
as considered earlier, but is just highly colour polymorphic. Additional notes on its morphology and
colouration are provided. Needs for further surveys in the Andaman Islands for documenting the
distribution of this species are highlighted.
INTRODUCTION
The Andaman and Nicobar Islands are situated south of Burmese peninsula, in the Bay of Bengal (India).
Though exploration of its fauna began way back in the mid 19th century (Blyth, 1846), anuran fauna of the Andaman
Islands is scantily explored and is currently known to comprise eight species belonging to six genera and three
families (Das, 1999). Fairly recent among the discoveries was the frog Rana charlesdarwini Das, 1998, which was
described from the forests of Mount Harriet National Park, South Andaman. Ever since its description, this species
has had an uncertain generic as well as family level placement. Das (1998) tentatively regarded it to be a member
of the family Ranidae and genus Rana Linnaeus, 1758 sensu Boulenger (1920). Dubois (in Dubois et al., 2005: 42)
stated that, having a forked omosternum, it was “clearly not a member of the genus Rana, let alone of the Raninae”,
and referred it “to the Limnonectini, without generic allocation”. Consequently, Das & Dutta (2007) referred it to
the genus Limnonectes Fitzinger, 1843 without any explicit reason. Frost et al. (2006) did not mention this species.
Frost (2015), without further explanation except the unclear formula “by implication”, transferred it to the genus
Ingerana Dubois, 1987 in the subfamily Occidozyginae of the family Dicroglossidae.
To this day, Ingerana charlesdarwini (Das, 1998) has been defined only based on three adult type specimens
and five tadpoles, all collected together from Mount Harriet National Park in South Andaman Island (Das, 1998).
Very little information is available on its distribution and natural history apart from the fact that it breeds and
lays eggs in water-filled tree holes (Das, 1998). Recent status assessments regard Ingerana charlesdarwini to
be Critically Endangered (Das et al., 2004) and it is also considered to be one among India’s “lost” amphibians
(Anonymous, 2011). This paper presents further information on intraspecific morphological variation in Ingerana
charlesdarwini based on fresh specimen collections.
Received 27 December 2015
*Corresponding author
Accepted 01 October 2016
Corresponding editor Annemarie Ohler
[email protected]
Published Online 17 March 2017
© ISSCA and authors 2017
S. R. CHANDRAMOULI
MATERIALS AND METHODS
This published work has been registered in ZooBank. The ZooBank LSIDs (Life Science Identifiers)
can be resolved and the associated information viewed through any standard web browser. The LSID for this
publication is: urn:lsid:zoobank.org:pub:4CCFC854-954C-4DB5-8173-12515B142EB3
Specimens were hand-collected, photographed in situ, fixed in 70 % ethanol, and are deposited in the
herpetological collection of the Wildlife Institute of India, Dehradun, India (WII). WII 45-46, WII 48-51, WII
76 and WII 104 were collected from primary evergreen forests in Mount Harriet National Park, South Andaman
Island (11.74º N, 92.73º E, 307 m a.s.l.) between December 2010 and March 2011. WII 12 and WII 73-74 were
collected from secondary forests in Long Island near Middle Andaman (12.37º N, 92.92º E, 58 m asl) during May
2010. The type specimens at the Zoological Survey of India (ZSI), Kolkata (ZSI A.8890-8892), collected from
Mount Harriet National Park, South Andaman, were re-examined. Geographic co-ordinates of the localities were
taken from Google EarthTM software.
The following morphometric measurements were recorded using MitutoyoTM dial calipers to the nearest
0.01 mm after three to four months of preservation: snout-vent length (SVL), measured from the tip of the snout
till the cloaca; head length (HL), measured from the tip of the snout to the posterior edge of the mandible; head
width (HW), measured at the jaw angle; horizontal diameter of the orbit (ED); distance between anterior border
of the eye to posterior edge of the nostril (EN); distance between anterior border of the eye to the snout tip (ES);
distance between the anterior margins of the upper eyelids (IO); closest distance between the nares (IN); horizontal
diameter of the tympanum (TYH); upper arm length (UAL), measured from the point of insertion of the forelimb
to the trunk to elbow; lower arm length (LAL), measured from the elbow to the base of the outer metacarpal
tubercle; palm length (PAL), measured from the posterior border of the outer metacarpal tubercle to the tip of
the third finger; femur length (FEL), measured from the cloaca to the tip of the knee; tibia (shank) length (TBL),
measured from the knee to the obtuse margin of the tibia; foot length (FOL), measured from the anterior end of
the tarsus to the tip of the fourth toe. Webbing formula follows Savage & Heyer (1997). Type specimens were
measured by the author in order to account for inter-observer bias in the measurements. A total of eleven specimens
of this species belonging to three different colour morphs were collected and are briefly described below. Colour
descriptions made here are based on the observation of live individuals and their photographs in life. The above
measurements were standardized to SVL and were subjected to principal component analysis with varimax axis
rotation, to examine the distribution of these different colour morphs in the multivariate morphological space.
TAXONOMY
Ingerana charlesdarwini (Das, 1998)
Rana charlesdarwini Das, 1998: 25.
Limnonectes charlesdarwini — Das & Dutta, 2007: 157.
Diagnosis. A small to medium
sized dicroglossid characterized by:
absence of lingual papilla; presence
of vomerine teeth; skin with minute,
non-uniform granules scattered all
over the dorsum; venter smooth; finger
and toe tips with swollen discs lacking
circum-marginal grooves; reduced toe
webbing, not exceeding the penultimate
subarticular tubercle on 4th toe; distinct
and exposed tympanum.
Description and variation.
Females (mean SVL 32.8 mm ± 2.28;
n = 8) slightly larger than males (mean
SVL 28.2 mm ± 1.67; n = 6); head
longer in males (mean HL/SVL 0.37 ±
0.04) than in females (mean HL/SVL
Figure 1. An adult male Ingerana charlesdarwini (uncollected) in life.
48
ALYTES 2017 | 33
0.35 ± 0.03), as broad as long in females (mean HL/HW 0.99 ± 0.09) and slightly broader than long in males
(mean HL/HW 0.96 ± 0.07). Eyes equally large in males and females, less than half the length of the head (mean
ED/HL 0.32 ± 0.04). Snout tip blunt and rounded in both dorsal and lateral views; vomerine teeth present; in two
rows well separated from each other; tongue large and bifid, lacking lingual papilla; nostrils located closer to the
snout tip than to the eyes; slightly more closer in males (EN/ES -0.66 ± 0.08) than in females (EN/ES -0.58 ± 0.02).
Tympanum smaller than the eye, relatively smaller in males (TYH/ED -0.6 ± 0.14) than in females (TYH/ED -0.71
± 0.08). Supra tympanic fold well developed, commencing from the posterior corner of the eye till the jaw angle
in the form of a thick, fleshy ridge. Upper arms short, less than a quarter of the body in both the sexes (mean UAL/
SVL 0.19 ± 0.02); thigh shorter in males (mean FEL/SVL 0.44 ± 0.04) than in females (mean FEL/SVL 0.47 ±
0.04) and slightly longer than the tibia in both sexes (mean FEL/TBL 0.89 ± 0.05); foot equally shorter than the
thigh in both sexes (mean FEL/FOL 0.97 ± 0.09). Under surface of the digits with well-developed subarticular
tubercles; tips swollen but not dilated; terminal discs lacking circum marginal grooves. Relative length of the
fingers: III = I > IV > II; toes: IV > III > V > II > I. Webbing absent in fingers, toes partially webbed: webbing
formula I½ – 1II½ – 2III1 – 2IV2 – 1V. See fig. 1 and more deatailed measurements on tab. 1-2.
HL
HW
ED
EN
ES
IO
IN
TYH
UAL
LAL
PAL
♀
31.9
10.5
11.59
3.57
2.58
4.69
2.69
3.12
2.51
6.22
6.57
7.83
T2
♂
28.32 11.12 11.04
3.18
2.5
3.98
2.37
2.65
1.96
4.25
5.42
6
WII 73
T2
♂
28.74
8.81
10.33
3.37
2.46
3.87
2.65
2.65
1.25
3.72
4.61
6.18
10.91 12.06 12.34
WII 74
T2
♂
27.98
9.49
10.45
3.38
2.64
4.16
2.22
2.52
2.11
4.46
4.87
6.27
12.46 13.99 12.91
WII 49
T2
♀
32.6
12.41 12.73
3.85
3.11
5.01
2.35
2.6
2.92
4.9
6.01
7.5
14.88 16.68
WII 50
T2
♀
32.84 10.91 11.01
4.22
2.92
4.9
2.59
2.83
2.67
6.65
7.51
8.28
17.45 18.58 16.89
WII 51
T1
♀
33.19 10.84 12.01
3.4
2.52
4.18
2.9
2.9
2.89
4.99
6.4
7.39
16.66
WII 46
T1
♀
32.06 13.67
11.4
3.22
3.95
4.92
2.37
3.0
2.5
6.32
6.69
7.27
15.77 16.27 15.27
WII 48
T3
♀
37.94 13.29 13.53
3.67
3.18
5.13
2.64
3.0
2.95
5.48
6.5
7.18
17.85 18.58 16.89
WII 45
T3
♂
24.54
8.82
3.49
2.29
3.82
1.67
2.05
1.8
3.87
5.39
5.17
11.47 13.54
WII 76
T3
♀
29.16 10.65 10.42
3.61
2.75
4.52
2.16
2.54
2.35
5.67
5.82
7.15
12.5
ZSI A.8890
T3
♂
27.39
10.64
3.31
2.45
3.83
1.89
2.49
2.44
4.98
5.63
5.79
13.46 14.83 12.42
ZSI A.8891
T2
♀
32.43 13.04 13.32
4.25
3.15
5.93
2.31
3.48
2.7
6.66
6.94
7.43
17.7
ZSI A.8892
T2
♀
32.34 11.16 11.91
3.72
2.98
5.56
2.6
3.02
2.78
7.03
7.33
6.92
15.42 18.47 15.06
8.77
10.7
FOL
SVL
T2
WII 12
TBL
Sex
WII 104
Voucher
No.
FEL
Morph
Table 1. Measurements of Ingerana charlesdarwini individuals.
13.03 17.62 16.57
11.7
12.68 12.07
17.8
15.8
16.1
12.7
14.67 15.25
18.51 16.04
Colouration in life. The colour patterns of the three different morphotypes observed during this study are
described below (see fig. 2):
Morphotype T1 (♀:WII 46, WII 51): Overall body colouration dull olive brown, with a bright orange
coloured ‘butterfly shaped’ marking on the dorsum situated posterior to the occiput, and a smaller spot of the same
colour above the sacral region. Black subocular spots very feeble and the limbs lacking barred pattern. Specimens
of this morph are referred in the analysis with a prefix “T1”.
Morphotype T2 (♀: ZSI A.8891-8892, WII 49-50, WII 104; ♂: WII 12, WII 73-74): Specimens of
this morph were observed to be able to change the intensity of body colouration depending on the amount of
environmental moisture conditions. This morph is light tan coloured with very distinct black bars on the limbs, a
feeble “W” shaped marking on the mid-dorsum with two indistinct orange spots. Occasionally, a mid-dorsal stripe
was observed on the body, extending from the snout-tip till the cloaca. In the presence of water, it turns dark brown
dorsally with two small, orange coloured spots, the larger one behind the occiput and the smaller one at mid-body.
Male have a dark gular vocal sac. Limbs are distinctly barred. Specimens of this morph are referred in the analysis
with the prefix “T2”.
Morphotype T3 (♀: WII 48, WII 76; ♂: WII 45, ZSI A.8890): This very distinct colour morph is bright
creamy white overall with a dark brown patch on the dorsum, with a very thin, creamy white mid dorsal stripe and
49
S. R. CHANDRAMOULI
Figure 2. Different colour morphs of Ingerana charlesdarwini observed during the study and the type series: A. (T1) WII 46 ♀ in life; B. (T2)
Paratype ZSI A.8891 ♀; C. (T2) WII 73 ♂ in life; D. (T3) ZSI A.8890 ♂ holotype; E. (T3) WII 76 ♀ in life; F. (T3) WII 48 ♀ showing mature
ova.
two distinct white subocular spots. This morph matchs well with the description of the male holotype. Male have
a dark gular vocal sac. Specimens of this morph are referred in the analysis with the prefix “T3”.
Multivariate morphometric analyses carried out using the above variables could not discern any differences
between the different colour morphs identified and also between the two sexes. Additionally, a lack of consistency
in the colour pattern was observed between the sexes (fig. 3).
On some overlooked historical specimens
Before the present work, Ingerana charlesdarwini (originally described as Rana charlesdarwini) was
known only from three adult type specimens. While examining the type series at the Zoological Survey of India,
Kolkata, a collection of specimens (ZSI 19525–27 and ZSI 19529) labelled as Rana doriae andamanensis collected
50
ALYTES 2017 | 33
Table 2. Significance of differences in measurement
by Nelson Annandale from “small streamlet running through
ratios between sexes of Ingerana charlesdarwini.
deep jungle, S. Andamans” and “thick jungle W. side of mount
Harriet S. Andaman” was also traced. The best preserved
specimen of the lot, “ZSI 19526?” (fig. 4) was examined
U
Z
p
and was found to exhibit the following morphological
0
-2.93
0.003
SVL
characteristics diagnostic of Ingerana charlesdarwini: (1)
HL:SVL
22
0
1
absence of lingual papilla; (2) presence of vomerine teeth;
(3) moderate degree of toe webbing, reaching the penultimate
HL:HW
22
0
1
subarticular tubercle of the 4th toe; and (4) finger tips with
UAL:SVL
11
-1.47
0.13
swollen discs lacking circum marginal grooves.
FEL:TBL
18
-0.53
0.59
Given the data on its provenance, this specimen with
ED:HL
15
0.93
0.34
the abovementioned characters can readily be differentiated
EN:ES
9
-1.74
0.08
from other geographically sympatric dicroglossid frogs
TYH:ED
4
-2.4
0.01
known from the Andaman Islands (Das, 1999; Chandramouli
FEL:SVL
11.5
-1.4
0.16
et al., 2015) as follows: dorsal colouration dark and indistinct
FEL:FOL
18
-0.53
0.59
from the lateral body (vs. unpatterned chestnut brown with
darker flanks in Fejervarya andamanensis; dull brown above
with dark spots on the dorsum with a “W” shaped marking
in Limnonectes cf. hascheanus); nostrils located laterally (vs.
dorsally in Fejervarya cf. cancrivora); toe webbing reduced, reaching only the penultimate subarticular tubercle on
toe IV (vs. greater degree of webbing, extending till the distal subarticular tubercle in Limnonectes cf. hascheanus
and Fejervarya cf. cancrivora); and presence of a small outer metatarsal tubercle (vs. absent in Fejervarya cf.
cancrivora). This specimen does not exhibit any non-diagnostic character compared to I. charlesdarwini, even
after this expanded morphological characterization. Hence, given the morphological congruence and locality data,
I hereby refer this specimen to the nominal species Ingerana charlesdarwini.
The jar label-name Rana doriae andamanensis associated with these specimens has not appeared anywhere
previously in published literature. Checklists
that include “Rana doriae” from Andaman
Islands - e.g. Das (1999), as Limnonectes
doriae (Boulenger, 1887) - did not mention
this
trinominal
representation.
The
Table 3. Factor loadings and % variance explained by
the principal components (maximum loadings for each
character in bold).
Figure 3. Plot of principal component analysis of specimens of the
different morphotypes of Ingerana charlesdarwini based on standardised
morphometric measurements. Squares – males (solid squares – morphotype
T3; white squares – morphotype T2); circles – females (solid circles –
morphotype T2; white circles – morphotype T3; red circles - morphotype
T1). Type specimens with a dot in the middle. Note the lack of separation
between the different colour morphs or the sexes.
PC 1
PC 2
HL
0.54
2.49
HW
-0.07
-1.55
ED
1.96
2.73
EN
-1.37
1.57
ES
1.51
-2.07
IO
-2.51
-0.99
IN
0.98
0.77
TYH
4.01
-1.47
UAL
1.98
0.49
LAL
-2.32
-1.78
PAL
-5.05
0.71
FEL
-1.29
0.00
TBL
0.95
0.63
FOL
0.67
-1.54
Eigenvalue
5.34
2.56
% variance
38.17
18.25
51
S. R. CHANDRAMOULI
synchronous recognition and mention of Limnonectes
andamanensis (Stolizcka, 1870) in such lists indicate
that Rana doriae andamanensis was not recognised
as an elevated, species-level taxon. Thus, being a
jar-label name, Rana doriae andamanensis, without
any explicit mention of taxon author(s) and year of
publication, is deemed to be an unavailable name in
the sense of the Articles 10-11 of the 4th edition of the
Code (1999). The only amphibian type catalogue of this
holding and its associated branch offices (Chanda et
al., 2000) did not mention these specimens anywhere,
indicating that the specimens were never considered
as types, contrary to Annandale’s purported intent.
DISCUSSION
Owing to the explicit but clearly incorrect
generic placement of this taxon in its original
description (Das, 1998), the “comparisons” made with
“related taxa” were very vague, irrelevant and dilute.
Figure 4. Voucher specimen ZSI 19526? labelled as Rana doriae
I hereby furnish a proper species-level comparison
andamanensis, collected by Nelson Annandale.
with the revised set of congeners. Brown et al. (2015)
recently reviewed and redefined the content and
distribution of the genus Ingerana Dubois, 1987.
Species formerly attributed to this genus from parts of
the Sundaland (i.e., mariae, rajae, baluensis and sariba) were transferred to a new Sundaic, ceratobatrachid genus
Alcalus Brown, Siler, Richards, Diesmos & Cannatella, 2015. Following this revision, Yan et al. (2016) studied
the phylogenetic position of the members xizangensis, medogensis and alpina that were previously attributed to
Ingerana, and reallocated them to the genus Liurana Dubois, 1987 under the family Ceratobatrachidae, extending
its distribution range to the Indo-Chinese region. These authors particularly speculated that the Andamanese
species charlesdarwini could also belong to a sister lineage to the ceratobatrachid frog genus Liurana. However,
Das (1998) specifically remarked that charlesdarwini differs from the genus Liurana in lacking a lingual papilla.
This situation makes it difficult to propose an accurate generic allocation for charlesdarwini. However, considering
the geographic proximity of charlesdarwini to the type species of genus Ingerana (I. tenasserimensis) and its
morphological attributes, the generic allocation proposed by Frost (2015) is followed herein provisionally until
finer taxonomic resolution is attained. Ingerana currently has four representatives distributed in parts of Indochina.
Ingerana charlesdarwini, one of the westernmost representatives of the genus, can be distinguished from congeners
based on the following combination of characters: presence of poorly developed discs lacking circum-marginal
grooves (vs. small discs with evident circum-marginal grooves in I. tenasserimensis); presence of an evidently
exposed tympanum (vs. indistinct tympanum, hidden under the skin in I. reticulata) and reduced toe-webbing, not
exceeding ante-penultimate sub-articular tubercle (vs. extensively webbed toes in Ingerana borealis) (Sailo et al.,
2009; Borah et al., 2013).
At species-level, the data presented here based on new collections add to the current knowledge on this
poorly known taxon. In the original description, it was considered to be sexually dimorphic and dichromatic,
with females being larger and “strikingly different” in colouration from the only known male (Das, 1998). The
analysis presented here based on a larger sample size suggests that the different morphotypes designated based
on their colouration fall well within the same morphological entity and are not separable into different groups
(fig. 3; tab. 3). Thus, the correlation between the body colouration and sex of the specimens assumed in the original
description should be attributed to an artefact due to a very low sample size. Also, two female specimens (T3 WII
45, WII 76) with their colouration exactly matching that of the male holotype (T3 ZSI A.8890), and four male
specimens (T2 WII 12, WII 49–50, WII 74) with their colouration matching the description of female paratypes
(ZSI A.8891–8892) but for the presence of dark blackish colouration in the gular vocal sac, clearly indicate that
this species is not sexually dichromatic as considered earlier (Das, 1998) but is highly colour polymorphic.
Apart from Mount Harriet National Park, Ingerana charlesdarwini was recorded from Long Island near
Middle Andaman. There have been no authentic records of this species after its description but for that by Das et al.
(2004) who included Saddle Peak, situated in North Andaman Island, in its distribution range, without any further
52
ALYTES 2017 | 33
information on the location or the individuals sighted. This situation calls for an assessment of its present status
and geographic range. Hence, further surveys throughout the Andaman Islands are necessary to assess the present
status and document the exact distribution range of this poorly known, Critically Endangered species.
ACKNOWLEDGEMENTS
I thank the Wildlife Institute of India for the award of Junior Research fellowship and supporting my
research; the Department of Science and Technology for funding; the staff of ZSI, Kolkata for the facilities
provided; Sudeer, Anant, Glen Xavier and Harikrishnan for their assistance with fieldwork.
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Yan, F., Jiang, K., Wang, K., Jin, J., Suwannapoom, C., Li, C., Vindum, J.C., Brown, R.M., Che, J. (2016). The
Australasian frog family Ceratobatrachidae in China, Myanmar and Thailand: the discovery of a new
Himalayan forest frog clade. Zoological Research 37(1): 7-14.
APPENDIX
List of specimens examined
Island.
WII 45–46, WII 48–51, WII 76 and WII 104: Mount Harriet National Park, South Andaman Island.
WII 12, WII 73–74: Long Island.
ZSI A.8890 (Holotype), ZSI A.8891–8892 (paratypes); Mount Harriet National Park, South Andaman
ZSI 19526?: labelled as Rana doriae andamanensis; “S. Andamans”.
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