A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species
Author
Pal, Saunak
Author
Vijayakumar, S. P.
Author
Shanker, Kartik
Author
Jayarajan, Aditi
Author
Deepak, V.
text
Zootaxa
2018
2018-09-19
4482
3
401
450
journal article
29419
10.11646/zootaxa.4482.3.1
5916d93e-11d4-49c5-8085-8bcd37d3739b
1175-5326
1440674
10258391-162F-4C7D-AA5E-1A03A4F3FE19
Calotes
Cuvier, 1817
The genus
Calotes
Cuvier, 1817
with the
type
species
Lacerta calotes
Linnaeus, 1758
belongs to the subfamily
Draconinae
of the family
Agamidae
, containing 26 species. The geographical distribution of this genus is largely restricted to
South
Asia and Indo-china, except for one isolated species
Calotes nigriplicatus
Hallermann, 2000
found in
Ambon
Island in
Indonesia
. The greatest diversity of the genus is in the
Western
Ghats and
Sri Lanka
. Four species of
Calotes
(
C. grandisquamis
,
C. nemoricola
,
Calotes
cf.
versicolor
,
Calotes calotes
) are known from the
Western
Ghats of which two are endemic to this region.
FIGURE 4.
Lateral photograph showing live coloration of A. adult female
C. calotes
and B. adult male
C.
cf. versicolor
.
Diagnosis.
Medium to large sized agamids with a relatively broader head; dorsal and lateral scales uniform, arranged in rows of backwardly pointed scales; dorsal and nuchal crest more or less developed; groups or rows of spines from eye to above tympanum; often an antehumeral skin fold or pit present covered with small scales; gular sac usually present; tympanum naked; tail long and slender, usually swollen and rounded at the base in males; no preanal or femoral pores.
FIGURE 5.
Lateral photograph showing live coloration of A. adult male
C. grandisquamis
and B. adult male
C. nemoricola
.
Distribution.
Lizards belonging to the Genus
Calotes
are widely distributed across
Iran
,
Afghanistan
,
Bhutan
,
Bangladesh
,
Maldives
,
Nepal
,
India
,
Pakistan
and
Sri Lanka
as well as southern
China
; parts of
Myanmar
,
Thailand
,
Laos
,
Cambodia
,
Vietnam
, Peninsular
Malaysia
and parts of
Indonesia
. Moreover,
Calotes versicolor
has been introduced into
Kenya
, Borneo,
Sulawesi
,
Seychelles
,
Mauritius
,
Oman
and
Florida
(
USA
) (Das
et al
. 2008;
Manthey, 2008
;
Uetz and Hošek, 2016
).
FIGURE 6.
Lateral photograph showing live coloration of A. adult male
Monilesaurus rouxii
comb. nov
.
and B. adult male
Monilesaurus ellioti
comb. nov
.
Taxonomic groups.
Smith (1935)
separated members of the genus
Calotes
into four distinct groups, I.
C. cristatella
group, II.
C. microlepis
group, III.
C. versicolor
group, and IV.
C. liocephalus
group, each composed of multiple species and another group composed of two dwarf species based on their morphological similarities. Later,
Moody (1980)
separated the genus into four different genera, namely
Bronchocela
Kaup, 1827
(group I of Smith),
Dendragama
Doria, 1888
,
Pseudocalotes
Fitzinger, 1843
(group II of Smith) and
Calotes
(group III, IV and the two dwarf species of Smith). Most of the
Calotes
species occurring in the
Western
Ghats belong to the
Calotes versicolor
group of
Smith (1935)
except
Calotes rouxii
and
Calotes ellioti
, which belong to the dwarf group. Here, we transfer these two species to the new genus
Monilesaurus
gen. nov
.
In the present study, we follow Smith’s species groups for taxonomic accounts and morphological comparisons. Since Smith, there have been various new additions to the genus
Calotes
and to the
Calotes versicolor
group. In this study, we compare each individual species only with members of their own group.
Calotes versicolor
group.
Members of the
C. versicolor
group are characterized by a medium to large adult body size; body more or less compressed, dorsal and lateral scales large, uniform, smooth or feebly keeled and oriented postero-dorsally; antehumeral fold absent; dorsal scales larger than the ventrals except in
C. calotes
where it is equal to or slightly smaller than the ventrals; nuchal and dorsal crest well developed, continuous, at least till behind the shoulders; postorbital spine absent except in
C. emma
Gray, 1845
; supratympanic spines in form of paired patches or as a single longitudinal series of spines (
Fig. 4
&
5
). For taxonomic comparisons, we add the recently designated
C. minor
to Smith’s
C. versicolor
group (See
Deepak
et al.
2015
).
FIGURE 7.
Lateral photograph showing live coloration of A. adult male
Monilesaurus acanthocephalus
gen. et sp. nov.
and B. adult male
Monilesaurus montanus
sp. nov.
Content.
Calotes bachae
Hartmann, Geissler, Poyarkov,
Ihlow, Galoyan, Rödder & Böhme, 2013
;
C. bhutanensis
Biswas, 1975
;
C. calotes
;
C. ceylonensis
Müller, 1887
;
C. chincollium
Vindum, 2003
;
C. desilvai
Bahir & Maduwage, 2005
;
C. emma
;
C. grandisquamis
,
C. hutunwini
Zug & Vindum, 2006
;
C. irawadi
Zug, Grown,
Schulte & Vindum, 2006
;
C. jerdoni
Günther, 1870
;
C. liocephalus
Günther, 1872
;
C. liolepis
Boulenger, 1885
;
C. versicolor
,
C. manamendrai
Amarasinghe & Karunarathna, 2014
;
C. maria
Gray, 1845
;
C. medgoensis
Zhao & Li, 1984
;
C. minor
,
C. mystaceus
Duméril &
Bibron, 1837
;
C. nemoricola
,
C. nigrilabris
Peters, 1860
;
C. nigriplicatus
Hallermann, 2000
and
C. pethiyagodai
Amarasinghe, Karunarathna, Hallermann, Fujinuma, Grillitsch & Campbell, 2014
.
Diagnosis.
All the species belonging to this group can be differentiated from the other groups of
Calotes
based on the combination of the following characters: From members of the genus
Monilesaurus
gen.nov.
, they can be easily distinguished morphologically based on a larger adult size (vs. smaller); dorsal and lateral scales rows directed posterodorsally, much larger in size (vs. posteroventrally, smaller in size); dorsal and lateral scales larger than ventrals, except in
C. calotes
(vs. not larger than ventrals) (
Fig. 4–5
vs 6–7) and absence of antehumeral fold (vs. presence of a well developed antehumeral fold) (
Fig. 10
). From members of the genus
Psammophilus
,
group they can be distinguished by the presence of compressed body shape (vs. depressed) (
Fig. 4–5
vs 8a); absence of an antehumeral fold (vs. antehumeral fold present) (
Fig. 10f
vs
10g
); lateral scales much larger (vs. smaller); presence of well developed nuchal crest (vs. reduced) and lesser number of midbody scale rows, less than 60 (vs. scale rows higher in number, more than 80). From
Microauris
gen. nov.
by having a relatively large tympanum (
Fig. 9a
vs 9b); scales on head large uniform shield like (vs. small, sub-triangular) (
Fig. 10b
vs 10d).