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Rediscovery of Calotes andamanensis BOULENGER, 1891, and assessment of its generic allocation

Authors:
Harikrishnan Surendran at Wildlife Trust of India
Harikrishnan Surendran
Karthikeyan Vasudevan at Centre for Cellular and Molecular Biology
Karthikeyan Vasudevan
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Abstract and Figures

wiederentdeckung von Calotes andamanensis Boulenger, 1891 und die Beurteilung seiner gattungszugehörigkeit (squamata: sauria: Agamidae) surenDrAn hArIKrIshnAn & KArThIKeyAn VAsuDeVAn KurZFAssung Die nur vom holotypus bekannte draconine Agame Calotes andamanensis Boulenger, 1891, wurde auf den Andamanen (India) wiedergefunden. Die Autoren legen morphometrische und meristische Daten von sechs exemplaren von drei Inseln des Andamanen-Archipels vor und machen ergänzende Angaben zur Beschreibung des Typusexemplares. nach der äußeren Morphologie der echsen ist ihre Zuordnung zur gattung Calotes nicht zutr-effend, vielmehr sollten sie in die gattung Pseudocalotes gestellt werden. ABsTrACT The little known draconine agamid lizard Calotes andamanensis Boulenger, 1891, was rediscovered from the Andaman Islands (India). The authors present morphometric and meristic data from six specimens originating from three islands of the Andaman Archipelago, and thereby add information to the description of the type speci-men. The lizard's external morphology suggests that its generic assignment to Calotes in its present definition can-not be maintained and that this species should better be included in the genus Pseudocalotes.
Metric data (mm) from six specimens of Pseudocalotes andamanensis (Boulenger, 1891) from the Andaman Islands studied in this paper. Data of the holotype (ZMuC 36944) was taken from KrIshnAn (2008). For definition of measured parameters see Materials and Methods. M – Male, F – Female. Tab. 1: Metrische Daten (mm) von sechs in dieser Arbeit untersuchten Pseudocalotes andamanensis (Boulenger, 1891) von den Andamanen Inseln. Die Daten des holotypus (ZMuC 36944) stammen aus KrIshnAn (2008). Die Meßstreckendefinitionen sind in Material und Methoden angegeben. M – Männchen, F – weibchen.
Metric data (mm) from six specimens of Pseudocalotes andamanensis (Boulenger, 1891) from the Andaman Islands studied in this paper. Data of the holotype (ZMuC 36944) was taken from KrIshnAn (2008). For definition of measured parameters see Materials and Methods. M – Male, F – Female. Tab. 1: Metrische Daten (mm) von sechs in dieser Arbeit untersuchten Pseudocalotes andamanensis (Boulenger, 1891) von den Andamanen Inseln. Die Daten des holotypus (ZMuC 36944) stammen aus KrIshnAn (2008). Die Meßstreckendefinitionen sind in Material und Methoden angegeben. M – Männchen, F – weibchen.
… 
Meristic data from six specimens of Pseudocalotes andamanensis (Boulenger, 1891) from the Andaman Islands compared with the holotype (ZMuC 36944). Data of the holotype was taken from Boulenger (1891) and KrIshnAn (2008). For definition of parameters see Materials and Methods. M – Male, F – Female. The counts on left and right sides of body are separated by a comma. Table 2: Meristische Daten von sechs in dieser Arbeit untersuchten Pseudocalotes andamanensis (Boulenger, 1891) von den Andamanen sowie die des holotypus (ZMuC 36944) im Vergleich. Daten des holotypus aus Boulenger (1891) and KrIshnAn (2008). Die Meßstreckendefinitionen sind in Material und Methoden angegeben. M – Männchen, F – weibchen. rechte und linke Zählwerte sind durch Kommas getrennt.
Meristic data from six specimens of Pseudocalotes andamanensis (Boulenger, 1891) from the Andaman Islands compared with the holotype (ZMuC 36944). Data of the holotype was taken from Boulenger (1891) and KrIshnAn (2008). For definition of parameters see Materials and Methods. M – Male, F – Female. The counts on left and right sides of body are separated by a comma. Table 2: Meristische Daten von sechs in dieser Arbeit untersuchten Pseudocalotes andamanensis (Boulenger, 1891) von den Andamanen sowie die des holotypus (ZMuC 36944) im Vergleich. Daten des holotypus aus Boulenger (1891) and KrIshnAn (2008). Die Meßstreckendefinitionen sind in Material und Methoden angegeben. M – Männchen, F – weibchen. rechte und linke Zählwerte sind durch Kommas getrennt.
… 
Pseudocalotes andamanensis (Boulenger, 1891) – enlarged and modified lamellae on the leading edge of the third toe (wII 126). Abb. 3: Pseudocalotes andamanensis (Boulenger, 1891) – Vergrößerte umgestaltete lamellen an der unterkante der dritten Zehe (wII 126).
Pseudocalotes andamanensis (Boulenger, 1891) – enlarged and modified lamellae on the leading edge of the third toe (wII 126). Abb. 3: Pseudocalotes andamanensis (Boulenger, 1891) – Vergrößerte umgestaltete lamellen an der unterkante der dritten Zehe (wII 126).
… 
Pseudocalotes andamanensis (Boulenger, 1891) males in life. A -wII 055; B -wII 056. Abb. 4: Pseudocalotes andamanensis (Boulenger, 1891). Männchen. A -wII 055; B -wII 056.
Pseudocalotes andamanensis (Boulenger, 1891) males in life. A -wII 055; B -wII 056. Abb. 4: Pseudocalotes andamanensis (Boulenger, 1891). Männchen. A -wII 055; B -wII 056.
… 
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All content in this area was uploaded by Harikrishnan Surendran
Content may be subject to copyright.
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Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 2
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;>GHI>C;G6A67>6AH
"C I=: ;DAADL>C< 9:H8G>EI>DC D; H>M
;G:H=AN 8DAA:8I:9 HE:8>B:CH I=: 8=6G68I:G
HI6I:H D; I=: =DADINE: D; C. andamanensis
6G: >C9>86I:9 >C E6G:CI=:H:H >; I=:N L:G:
9>;;:G:CI+E:8>B:CCJB7:GH<>K:C6G:I=:
;>:A986I6AD<J:CJB7:GH>CI=:8DAA:8I>DCD;
I=:/>A9A>;:"CHI>IJI:D; "C9>6 :=G6JC
,=:H: HE:8>B:CH 6G: E6GI D; 6C DC<D>C<
HIJ9N 6C9 L>AA 7: 9:EDH>I:9 >C I=: 2DD
AD<>86A+JGK:ND;"C9>6$DA@6I62+"8DA
A:8I>DCDC8DBEA:I>DCD;I=:HIJ9NCDI=:G
HE:8>B:C6B6A:;DJC9>C/6C9DDG+DJI=
C96B6C"HA6C9>C#6CJ6GNL6H8DA
A:8I:96C99:EDH>I:9>CI=:8DAA:8I>DCD;I=:
2DDAD<>86A+JGK:ND;"C9>6&6G>C:*:<>DC
6A+I6I>DC)DGIA6>GC96B6C6C9'>8D76G
"HA6C9H "C9>6 2+"  ,=>H L6H I=:
;>GHIHE:8>B:CD;I=>HHE:8>:H8DAA:8I:97NI=:
6JI=DGHD;I=>HE6E:G+J7H:FJ:CI:;;DGIHID
AD86I:I=>HHE:8>B:CL:G:JCHJ88:HH;JA
*+-%,+
Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 3
+!
*"$*"+!''
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Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 4
*:9>H8DK:GND;Calotes andamanensis (-%' *
555555555555555555555555555555555555555
><HDEEDH>I:E6<:77<:<:CV7:GA>:<:C9:+:>I:
><Pseudocalotes andamanensis (-%' * Y%6I:G6AK>:LD;=:69
2&-=DADINE:/""
77Pseudocalotes andamanensis (-%' * Y$DE;KDCA6I:G6A
2&-!DADINEJH/""
><Pseudocalotes andamanensis (-%' *YDGH6AK>:LD;=:69
2&-=DADINE:/""
77Pseudocalotes andamanensis (-%' *YDGH6A6CH>8=I9:H$DE;:H
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-%' *
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I=:C96B6C"HA6C9HHIJ9>:9>CI=>HE6E:G6I6D;I=:=DADINE:2&-L6HI6@:C;GDB$*"+!'' 
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>:&:WHIG:8@:C9:;>C>I>DC:CH>C9>C&6I:G>6AJC9&:I=D9:C6C<:<:7:C&Y&SCC8=:CY/:>78=:C
)6G6B:I:G /"" /"" /"" /"" /"" /"" 2&-
+:M && & & &
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,%      
,/      
!%       
!/       
!       
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#%      
(       
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,G%       
,G/      
,G!      
!JB       
*69       
      
      
      
      
      
:B       
,>7       
,      
,      
,      
,      
,      
%%      
7:ADLJEE:GA>EL=>I:L>I=6L=>I:HIG>E:
:MI:C9>C<ID7:=>C9INBE6CJB7D9NL>I=
L=>I: HEDIH 6C9 K:GI>86A 76GH I6>A 76C9:9
L>I=E6A:<G::C6C9L=>I:I=:<G::C8DADG
G6E>9AN 9>H6EE:6GH L=:C HIG:HH:9 6C9 I=:
L=DA: 6C>B6A IJGCH 7GDLC DC 86EIJG:
JA6G EDJ8= L=>I: DG N:AADL >C B6A:H
67H:CI>C;:B6A:H,LDD;I=:B6A:H;DJC9
;><=I>C< :68= DI=:G =69 I=: >CC:G E6GIH D;
A>EH6C9IDC<J:8DADG:97G><=IN:AADLAA
HE:8>B:CH IJGC:9 E6A: 7AJ>H=L=>I: DG
7GDLC>C6A8D=DA
,=>H HE:8>:H 6EE:6GH ID 7: 6G7DG:6A
EG:;:GG>C<I=:8GDLCDG86CDEND;IG::HAA
>C9>K>9J6AHL:G:H><=I:99JG>C<I=:G:A6I>K:
AN9G>:GH:6HDC7:IL::C#6CJ6GN6C9EG>A
Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 5
+!
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:><=I BDG: >C9>K>9J6AH D; I=>H INE: L:G:
D7H:GK:96AADC=><=7G6C8=:HD;IG::HJE
IDB67DK:<GDJC9(CI=:<GDJC9I=:N
BDK:98AJBH>AN6C9H::B:9ID7:G:A6I>K:AN
HADLBDK>C<>CI=:IG::H6AHD'DI=>C<:AH:
>H@CDLC67DJII=::8DAD<ND;I=>HHE:8>:H
DBB:CIHDCI=:INE:AD86A>IN
D;Calotes andamanensis
88DG9>C< ID I=: DG><>C6A 9:H8G>EI>DC
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manensis L6H 8DAA:8I:9 DC I=: C96B6C
"HA6C9H7NG:9:G>8@9DAE= *()+,(*
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L6HEDHI:9>CI=:'>8D76G"HA6C9H6C9EGD7
67ANC:K:GK>H>I:9I=:C96B6C"HA6C9H6C9
I=JHHE:8JA6I:9I=6II=:INE:EGD767ANDG><
>C6I:9 >C I=: '>8D76G 6C9 CDI >C I=:
C96B6C"HA6C9H,=>H>H=DL:K:G8DCIG6
9>8I:9 7N
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G:EDGI:9=>H>CI:G68I>DCHL>I=E:DEA:D;I=:
IG>7: \ G:6I C96B6C:H:] C:6G )DGI A6>G
+DJI= C96B6C "HA6C9 *()+,(*
 AI=DJ<=
*(
)+,(*
9>9 CDI
:A67DG6I: DC I=: C6IJG6A =>HIDGN D; I=:H:
>HA6C9HI=:6JI=DGH7:A>:K:I=6IB:CI>DC>C<
I=:67DK:>CI:G68I>DC>HHJ;;>8>:CI:K>9:C8:
;DG  *()+,(* =6K>C< K>H>I:9 I=:
C96B6C"HA6C9HAHD6E6GI;GDBI=:INE:
D;C. andamanensisI=:G::M>HIH6CJB7:G
D;DI=:G=:GE:IDAD<>86AHE:8>B:CH>CI=:8DA
A:8I>DC D; I=: 2DDAD<>H@ &JH:JB
-C>K:GH>IND;DE:C=6<:C8DAA:8I:97N
*()+,(*6C9A67:A:96H7:>C<;GDBI=:
C96B6C"HA6C9H,=:H:>C8AJ9:Laticauda
colubrina +!'"*Ophiophagus
hannah ',(*  Naja sagittifera
/%% A67:A:9Naja kaouthia %++('
 ;DG HNCDCNBN 8DBE /R+,*
34Bungarus andamanensis "+/+
+'1%  AD86A>IN 96I6 H6NH ZC96
B6C +DJI=[ EG:HJB67AN +DJI= C96B6C
"HA6C9 Trimeresurus andersonii ,!(
% A67:A:9Trimeresurus purpureo -
maculatus *1  D; L=>8= I=:
C96B6CI6MDCL6HEG:K>DJHAN8DCH>9:G:9
6HJ7HE:8>:H 8DBE &%!(,*  ,!(*)
34 Dendrelaphis pictus C96B6C
"HA6C9H EDEJA6I>DC CDL G:;:GG:9 ID 6H
Dendrelaphis andamanensis '*+('
3.( % .' *(("#' 4,=:
67DK: B6I:G>6A >H 8A:6G :K>9:C8: I=6I 
*()+,(* 9>9 >C9::9 EGD8JG: HE:8>B:CH
8DAA:8I:9 >C I=: C96B6C "HA6C9H ,=:G:
;DG:I=:INE:AD86A>IN <>K:C 6H ZC96B6C
"HA6C9H[ >H G:<6G9:9 ID 7: 8DGG:8I 7N I=:
6JI=DGHD;I=>HE6E:G
:C:G>86AAD86I>DCD;(-%' *]H
Calotes andamanensis
:H8G>7>C< Calotes andamanensis
(-%' *  B:CI>DC:9 I=6I >I BDHI
8ADH:AN G:H:B7A:9 Calotes liolepis R',
!*6HE:8>:H;DJC9>C+G>%6C@67JI
L>I=DJI HE:8>;N>C< L=N =: I=DJ<=I HD
+&",! ;DAADL:9(-%' *]HDE>C
>DC6C9EA68:9C. andamanensis >C=>H\lio-
cephalus <GDJE] ,=>H 8DC8:EI D; Calotes
=6H 8=6C<:9 H>C8: 6C9 B:B7:GH D; I=:H:
><Pseudocalotes andamanensis (-%' *
YCA6G<:96C9BD9>;>:9A6B:AA6:DCI=:
A:69>C<:9<:D;I=:I=>G9ID:/""
77Pseudocalotes andamanensis (-%' *
Y.:G<GUW:GI:JB<:HI6AI:I:%6B:AA:C6C9:G
-CI:G@6CI:9:G9G>II:C2:=:/""
Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 6
<GDJEH 6G: CDL 6HH><C:9 ID I=: <:C:G6
Bronchocela $-)  Pseudocalotes
",2"' *6C9Calotes -."*
&((1  !%%*&''  Q!&
 &!('1  "C I=:>G G:K>:L D;
Pseudocalotes !%%*&''  Q!&
 ED>CI:9 DJI BDGE=DAD<>86A 8=6G68
I:GH ID H:E6G6I: I=: <:C:G6 Pseudocalotes
Calotes 6C9Bronchocela";I=>H8A6HH>;>86
I>DC >H 688:EI:9 Calotes andamanensis >H
8A:6GANCDI6B:B7:GD;I=:<:CJHCalotes
-."*9J:IDI=:A68@D;EDHIDG7>I6A
6C9HJEG6INBE6C>8HE>C:H6C9I=:EG:H:C8:
D;6=:I:GD<:CDJH7D9NH86A6I>DC6G:A6I>K:
AN ADC< 6C9 C6GGDL =:69 6C9 6 GDL D;
:CA6G<:9H86A:H7:IL::CDG7>I6C9HJEG6A67>
6AH&!('1 9>H8JHH:9I=:>BEDG
I6C8:D;8:GI6>CBDGE=DAD<>86A8=6G68I:GH>C
I=: HNHI:B6I>8H D; 9G68DC>C: 6<6B>9H 6C9
ED>CI:9 DJI I=6I I=: HJ7D8JA6G GDL D;
:CA6G<:9 H86A:H >H 8=6G68I:G>HI>8 D; H:K:G6A
>CHJA6G<:C:G66HL:AA6HSalea *1
Andamanensis >H8:GI6>CANCDI6B:B7:GD;
I=:<:CJHSalea 6H>IA68@H>IH8DCHE>8JDJH
CJ8=6A 9DGH6A 6C9 86J96A 8G:HIH 6C9 :C
A6G<:9 @::A:9 9DGH6A 6C9 A6I:G6A H86A:H
+&",! &!('1 ,=:67H:C8:
D; 6 ADC<>IJ9>C6AH@>C ;DA9:MI:C9>C< ;GDB
I=:6C<A:D;I=:?6LIDI=:H=DJA9:GK:CIG6AH
I=6I 6G: HB6AA:G I=6C 9DGH6AH 6C9 I=: G:A6
I>K:ANH=DGII6>A8A:6GAN>C9>86I:HI=6Ianda -
manensis >H CDI 6 B:B7:G D; I=: <:CJH
Bronchocela 6H9:;>C:97N!%%*&''
Q!& 6C9 !%%*&'' 
"C H=DL>C< 6 GDL D; :CA6G<:9 H86A:H
7:IL::C DG7>I 6C9 INBE6CJB =:I:GD<:
C:DJH 9DGH6A H86A:H :CA6G<:9 IG>6C<JA6G
A6B:AA6:JC9:GI=:I=>G9ID:6C9A68@>C<6C
:CA6G<:9I6>A 76H:>C B6A:H I=>H HE:8>:H >H
BDHIH>B>A6GIDHE:8>:HD;I=:<:CJHPseudo -
calotesL=>8=6G:L>9:HEG:69>C+DJI=:6HI
H>6&((1 !%%*&'' Q!
& !%%*&'' & -"* 
!%%*&'' :I 6A  &!('1 
"C=6K>C<G:A6I>K:ANHA:C9:G6C9ADC<A>B7H
I=: EG:H:CI HE:8>B:CH D; I=: C96B6C
"HA6C9H 6G: BDHI H>B>A6G ID Pseudocalotes
tympanistriga *1 ;GDB #6K6 6C9
*:9>H8DK:GND;Calotes andamanensis (-%' *
,67A:&:G>HI>896I6;GDBH>MHE:8>B:CHD;Pseudocalotes andamanensis (-%' *;GDBI=:
C96B6C"HA6C9H8DBE6G:9L>I=I=:=DADINE:2&-6I6D;I=:=DADINE:L6HI6@:C;GDB(
-%' *
6C9$*"+!'' DG9:;>C>I>DCD;E6G6B:I:GHH::&6I:G>6AH6C9&:I=D9H&Y&6A:Y:B6A:
,=:8DJCIHDCA:;I6C9G><=IH>9:HD;7D9N6G:H:E6G6I:97N68DBB6
,67A:  &:G>HI>H8=: 6I:C KDC H:8=H >C 9>:H:G G7:>I JCI:GHJ8=I:C Pseudocalotes andamanensis
(-%' *  KDC 9:CC96B6C:C HDL>: 9>: 9:H !DADINEJH 2&-  >B .:G<A:>8= 6I:C 9:H
!DADINEJH 6JH (-%' *  6C9 $*"+!''  >: &:WHIG:8@:C9:;>C>I>DC:C H>C9 >C &6I:G>6A JC9
&:I=D9:C6C<:<:7:C&Y&SCC8=:CY/:>78=:C*:8=I:JC9A>C@:2S=AL:GI:H>C99JG8=$DBB6H<:IG:CCI
)6G6B:I:G /"" /"" /"" /"" /"" /"" 2&-
+:M &&& & &
+JEG6A67>6AH       
"C;G6A67>6AH       
"CI:GC6H6AH  
6CI=6AH   
+JEG68>A>6G>:H   
"CI:GDG7>I6AH 
)DHIB:CI6AH 
-EE:G=:69H86A:H $::A:9 $::A:9 $::A:9 $::A:9 $::A:9 $::A:9 $::A:9
-EE:GHCDJIH86A:H +BDDI= +BDDI= +BDDI= +BDDI= +BDDI= +BDDI=
CA6G<:9H86A:H

DCI:BEA:
+86A:H6GDJC9
      

B>97D9N
'J8=6AHE>C:H       
%6B:AA6:JC9:G
      
G9;>C<:G
%6B:AA6:JC9:G
      
I=ID:
CA6G<:9@::A:9
 
H86A:HDCI=><=
 ,=:DG><>C6AH86A:8DJCI<>K:C7N(-%' * L6H!DL:K:G+&",! 6C9$*"+!''
G:EDGI:9
Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 7
+!
*"$*"+!''
$.
+-.'
><Pseudocalotes andamanensis (-%' *B6A:H>CA>;:/""/""
77Pseudocalotes andamanensis (-%' *&SCC8=:C/""/""
Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 8
*:9>H8DK:GND;Calotes andamanensis (-%' *
+JB6IG6 L=:G:6H >I 6EE:6GH 8ADH:HI ID P.
microlepis (-%' *  ;GDB I=:
"C9D=>C:H: ):C>CHJA6 6C9 P. larutensis
!%%*&'' & -"* ;GDBL:HI
:GC&6A6NH>6>CB>97D9NH86A:8DJCIH6C9
H86A6I>DC!
%%*&''
& -
"*

GDB P. tympanistriga P. andamanensis
9>;;:GH>C=6K>C<6L>9:G=:69!/!%
>CP. andamanensis>CP. tympanistri-
gaADC<:G=>C9A>B7HD;+.%>C
P. andamanensis>CP. tympanistriga
HJEG6A67>6AH6C9>C;G6A67>6AH
 HJEG6A67>6AH 6C9  >C;G6A67>6AH >C P.
tympanistriga"C=6K>C<:CA6G<:9IG>6C<J
A6GA6B:AA6:DCI=:A:69>C<:9<:D;I=:I=>G9
ID: I=>H HE:8>:H >H H>B>A6G ID I=: "C9D
=>C:H:HE:8>:H<GDJED;Pseudocalotes
"I H=DJA9 7: G:B:B7:G:9 I=6I BDHI
9G68DC>C:6<6B>9<:C:G66G:9:;>C:9HDA:AN
76H:9 DC E=:C:I>8 8=6G68I:GH "C H:K:G6A
IG6>IHI=:HE:8>:H;GDBI=:C96B6CH9>;;:GH
;GDBDI=:GPseudocalotesBDHICDI67AN>IH
EGDEDGI>DC6I:ANADC<:GA>B7H(CANP. tym-
panistriga >H @CDLC ID =6K: H>B>A6G A>B7
EGDEDGI>DCH "C 699>I>DC BDHI HE:8>:H D;
Pseudocalotes:M8:EIP. saravacensis "' *
+,-"' 6G:>C=67>I6CIHD;BDJC
I6>C;DG:HIH L=:G:6H I=:C96B6C:H: HE:
8>:H>H@CDLC;GDBH:6A:K:AID67DJIB
6HA &DG:DK:G >I :M=>7>IH 6 L:6@ 6CI:
=JB:G6A;DA9L=>8=>HDI=:GL>H:EG:H:CI>C
<:C:G6HJ8=6HCalotes -."*HDB:
HE:8>:H Psammophilus ",2"' * 
Gonocephalus $-)Japalura *1
Acanthosaura *1:I8!DL
:K:G>CI=:6JI=DGH]K>:LI=:H:9>;;:G:C8:H
6G:CDIHJ;;>8>:CIANEGDCDJC8:9ID?JHI>;NI=:
G:8D<C>I>DC D; 6 C:L <:CJH ;DG I=:C96
B6C:H:HE:8>:H"CI=:67H:C8:D;6BDA:8
JA6GE=NAD<:CN6C976H:9DC:MI:GC6ABDG
E=DAD<N6ADC:>I>HBDHI6EEGDEG>6I:ID8DC
H>9:GI=>HHE:8>:H6H6B:B7:GD;I=:<:CJH
Pseudo calotes ",2"' *L=>8=>H6
=><=AN 9>K:GH: <:CJH 8DBEG>H>C< ;DJGI::C
HE:8>:H6C9H=DL>C<6L>9:6GG6ND;8=6G68
I:GH
Pseudocalotes andamanensis
(-%' *
C:L8DB7>C6I>DC
=G:HDCNBHCalotes andamanensis -
(-%' *  +&",!  $*"+!
'' 
>6<CDH>HG:A6I>K:ANADC<=:69!/
!%  ADC<>IJ9>C6AH86A: GDLH
6GDJC9 B>97D9N 9DGH6AH 6C9 A6I:G6AH
HBDDI=HDB:I>B:HL:6@AN@::A:9C:6GI=:
H68G6AG:<>DCK:CIG6AHHIGDC<AN@::A:99DG
H6AHD; E6G6K:GI:7G6A ADC<>IJ9>C6A GDLH
A6G<:G I=6C A6I:G6AH D; >GG:<JA6G H=6E:
ED>CI>C< EDHI:GD9DGH6AAN A6I:G6AH ED>CI>C<
EDHI:GDK:CIG6AAN A6I:G6AH 6C9 K:CIG6AH D;
H>B>A6G H>O: K:CIG6AH HA><=IAN >GG:<JA6G 6
GDLD;:CA6G<:9H86A:H7:IL::CHJEG6A67>6AH
6C9 DG7>I 7DG9:G:9 7N DC: DG ILD HB6AA:G
H86A: GDLH <JA6G H86A:H HB6AA:G I=6C K:C
IG6AHL:6@AN@::A:9<JA6GEDJ8=EG:H:CI>C
B6A:H 6CI:=JB:G6A ;DA9E>I L:6@AN 9:K:A
DE:9 CJ8=6A 8G:HI 8DBEDH:9 D; 
A6C8:DA6I:HE>C:H9DGH6A8G:HI69:CI>8JA6I:
G>9<: :CA6G<:9 8DC>86A A6B:AA6: JC9:G I=:
A:69>C< :9<: D; I=>G9 ID:   A6B:AA6:
JC9:G;DJGI=ID:=>C9A>B7A:C<I=
D; +.% I6>A A:C<I=   D; +.%
HA><=IAN8DBEG:HH:96II=:76H:
>HIG>7JI>DC=67>IH6C9=67>I6I
Pseudocalotes andamanensis (-
%' *>H6C6G7DG:6AHE:8>:HJHJ6A
AN ;DJC9 DC I=: IGJC@ D; IG::H C:6G I=:
8GDLC DG DC 7G6C8=:H ,=>H HE:8>:H >H
@CDLC ;GDB +DJI= C96B6C "HA6C9
*JIA6C9"HA6C96C9%DC<"HA6C9(L>C<ID
>IH6G7DG:6A=67>IH>I>HG6G:ANH><=I:96C9>H
A>@:AN ID D88JG >C BDG: >HA6C9H >C I=:
C96B6C "HA6C9H ,=: DCAN DI=:G C6I>K:
6<6B>9HE:8>:HI=6ID88JG>CI=:H6B:=67>
I6I6G:HE:8>:HD;I=:<:CJHCoryphophylax
",2"' * >C +,"'!'*  L=>8=
6G:EG>B6G>AN>C=67>I>C<I=:JC9:GHIDGN6C9
:6H>AN9>;;:G:CI>6I:9;GDBP. andamanensis
7NI=:HIGDC<IG6CHK:GH:H@>C;DA968GDHHI=:
H=DJA9:G 6C9 C:8@ Calotes 8; versicolor
-"'  EGD767AN 6C >CIGD9J8:9
HE:8>:H>CC96B6C"HA6C9HD88JGH>C'DGI=
C96B6C6C9I=:=6G7DGIDLCD;)DGIA6>G
>C +DJI= C96B6C 7JI >H G:HIG>8I:9 ID
6CI=GDED<:C>8BD9>;>:9A6C9H86E:H6C9CDI
;DJC9 >C ;DG:HIH Pseudocalotes andama-
nensis >H:6H>AN9>HI>C<J>H=:9;GDBC. versi-
color 7N I=: G:<JA6GAN 6GG6C<:9 HIGDC<AN
@::A:9 EDHI:GD9DGH6AAN 9>G:8I:9 7D9N
H86A:H I=: G:A6I>K:AN H=DGI =:69 6C9 ADC<
CJ8=6A6C99DGH6AHE>C:H>CI=:A6II:G
,=:=:GE:ID;6JC6D;C96B6C"HA6C9H
>H @CDLC ID =6K: 6;;>C>I>:H IDL6G9H I=:
Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 9
%*(&$*'+*.(*"+!$
*',!1+%%"'++% )=NAD<:CN
D; Cerberus +:GE:CI:H !DB6ADEH>C6: 6C9 E=NAD
<:D<G6E=ND;Cerberus rynchops9>K:GH>;>86I>DCD;6
8D6HI6A B6G>C: HC6@: >C +DJI=:6HI H>6 #DJGC6A D;
>D<:D<G6E=N(M;DG9
(-%' *    (C C:L DG A>IIA:
@CDLC"C9>6C6C9 &6A6N6CG:EI>A:H 6C976IG68=>6CH
CC6AH6C9&6<6O>C:D;'6IJG6A!>HIDGN%DC9DC

+">D<:D<G6E=ND;I=:6BE=>7>6CH
6C9 G:EI>A:H D; I=: C96B6C 6C9 '>8D76G "HA6C9H
"C9>6EE"C(,!9,GDE>86A>HA6C9=:G
E:ID;6JC6(G><>C8JGG:CI9>K:GH>IN6C98JGG:CIHI6IJH
BHI:G96BAH:K>:G
 *()+,(*    .D867JA6GN D;
9>6A:8IHHED@:C>CI=:'>8D76G6C9C96B6C"HA:HL>I=
6H=DGI688DJCID;I=:C6I>K:HI=:>G8JHIDBH6C9=67>IH
6C9 D; EG:K>DJH 6II:BEIH 6I 8DADC>O6I>DC 6A8JII6
(;;>8:D;I=:+JE:G>CI:C9:CID; DK:GCB:CI)G>CI>C<
EE
!%%*&'' #I6MDCDB>8G:K>:L
D; I=: <:CJH Bronchocela +FJ6B6I6 <6B>96:
L>I=I=:9:H8G>EI>DCD;6C:LHE:8>:H;GDB.>:IC6B
*JHH>6C #DJGC6A D; !:GE:IDAD<N &DH8DL  

!%%*&''#Q!&/G:K>:L
D; I=: <:CJH Pseudocalotes +FJ6B6I6 <6B>96:
L>I=9:H8G>EI>DCD;6C:LHE:8>:H;GDB/:HI&6A6NH>6
BE=>7>6*:EI>A>6%:>9:C
!%%*&'' #  & -"* # 
C:LHE:8>:HD; Pseudocalotes +FJ6B6I6<6B>96:
;GDB J@>I %6GJI /:HI &6A6NH>6 !:GE:IDAD<>86
%6LG:C8:
"+!/*'&+ "*:9>H8DK:GND;
Calotes andamanensis (-%' *  6C9 6 G:
6HH:HHB:CID;I=:INE:AD86A>IN#DJGC6AD;I=:DB76N
'6IJG6A!>HIDGN+D8>:IN&JB76>
$*"+!'' +  ':L HE:8>:H D; Calotes
*:EI>A>6 +FJ6B6I6 <6B>96: ;GDB I=: HDJI=:GC
/:HI:GC =6IH"C9>6#DJGC6AD;!:GE:IDAD<N!DJHIDC
:I8
&!('1 ++NHI:B6I>86C9I6MDBDC>8
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Calotes kingdonwardi +&",!  Japalura kaul -
backi +&",!  Salea kakhienensis '*+('
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E>IK>E:GH<:CJHTrimeresurus;GDBBDA:8JA6GHIJ9>:H
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9DCA6G:C9DC)G:HH
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+&",!&,=:;6JC6D;G>I>H="C9>6
>C8AJ9>C<:NADC6C9JGB6*:EI>A>66C9BE=>7>6
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869:B>8H/>A9A>;:"CHI>IJI:D;"C9>6;DGI=:DEEDGIJ
C>IN ID 8DC9J8I G:H:6G8= 6I I=: />A9A>;: "CHI>IJI: D;
"C9>6:=G6JC(JGH>C8:G:I=6C@HIDI=::E6GI
B:CI D; DG:HIH 6C9/>A9A>;:C96B6C 6C9 '>8D76G
"HA6C9H ;DG E:GB>HH>DC ID 8DC9J8I ;>:A9LDG@ >C I=:
C96B6C 6C9 '>8D76G "HA6C9H /: 6AHD I=6C@ I=:
:E6GIB:CI D;+8>:C8:6C9 ,:8=CDAD<N+, DK
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'D+*+(+/:I=6C@&D<:CHC9:GH:C
2DDAD<>86A &JH:JB -C>K:GH>IN D; DE:C=6<:C ;DG
EGDK>9>C<E=DID<G6E=HD;I=:=DADINE:D;Calotes an -
damanensis/:6AHDI=6C@I=:>G:8IDGD;I=:2DDAD
<>86A +JGK:N D; "C9>6 G  !  &JGI=N 6C9 G
$6JH=>@:JI>;DG;68>A>I6I>C<I=::M6B>C6I>DCD;HE:8
>B:CH >CI=:8DAA:8I>DCD; I=: 2+" $DA@6I6 /: 6G:
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I=6C@H 6G: 9J: ID + * =6C9G6BDJA>C6C9 #6B:H
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>CI=:C96B6C"HA6C9H
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C:AI=6IH:E6G6I:HI=:C96B6C"HA6C9H;GDB
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Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 10
*:9>H8DK:GND;Calotes andamanensis (-%' * 
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 KDC /R%%*+,(*-*"* 2DDAD<>H8=:G ,=:>A
GHI:G6C9*:EI>A>:C/>:C :GDA9H+D=CEE
EAH
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JI>DCHID6G:K>:LD;I=:Dendrelaphis pictus &%"'
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&'!' Y &JHTJB '6I>DC6A 9]!>HID>G:
C6IJG:AA: )6G>H G6C8: /"" Y />A9A>;:
"CHI>IJI: D; "C9>6 .:GI:7G6I: DAA:8I>DCH
:=G6 JC -II6G6@=6C9 "C9>6 2+" Y
2DDAD<>86A +JGK:N D; "C9>6 &6G>C:
*:<>DC6A+I6I>DC)DGIA6>GC96B6C6C9
'>8D76G"HA6C9H"C9>62+"Y2DDAD<>86A
+JGK:ND;"C9>6$DA@6I6"C9>6
Bronchocela cristatella $-!% 
Y 2+"  C:6G ,6KDN 2+" 
+$'6G'>8D76G
Bronchocela danieli ,"/*"  "+
/+ Y2+"=DADINE:6BE
7:AA6N G:6I'>8D76GC96B6C'>8D
76G "HA6C9H "C9>6 2+" +$%'
%>IIA:'>8D76G
Bronchocela rubrigularis !%%*
&''  Y 2+" +$' '6C
8DLGN
Calotes aurantolabium $*"+!''
 Y'!&=DADINE:$6A6@@69
&JC96CI=JG6> ,><:G *:H:GK: ,6B>A '69J
"C9>6
Calotes jerdoni -',!*  Y
2+"1JCC6CHDJI=:GC=>C6
Calotes mystaceus -&P*"%  "
*(' Y2+"&6C>EJG"C9>6
Calotes versicolor -"' Y
2+"HE:8>B:CH;GDB=>ED'DGI=
C96B6C 2+"   HE:8>B:CH ;GDB
=>ED'DGI=C96B6C
Pseudocalotes andamanensis (-
%' *  Y /""  %DC< "HA6C9
C96B6C  '>8D76G "HA6C9H "C9>6 /""
 /""  /""  /""  +DJI=
C96B6C "HA6C9 C96B6C '>8D76G
"HA6C9H "C9>6 /""  *JIA6C9 "HA6C9
C96B6C'>8D76G"HA6C9H"C9>6
Pseudocalotes austeniana '''
%  Y =DADINE: 2+"  =>AAH
C:6G!6GB6II>HH6B
Pseudocalotes microlepis (-%'
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Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 11
... The correct generic placement of the species first described by Boulenger in 1891 as Calotes andamanensis has long been uncertain. Harikrishnan and Vasudevan (2013) analysed the type specimen and others and reassigned the species to the genus Pseudocalotes Fitzinger, 1843 on the basis of morphology and obvious similarities between the species. The species within the genus Pseudocalotes, had until 1980 been generally placed within Calotes Daudin, 1802, whereupon Moody (1980 resurrected Pseudocalotes and transferred a number of species to the genus. ...
... This genus Notacalotes gen. nov. is alternatively defined as follows: relatively long head (hw: hl = 0.59); 56-67 longitudinal scale rows around midbody; dorsals and laterals smooth, sometimes weakly keeled near the sacral region; ventrals strongly keeled; dorsals of 4-7 paravertebral longitudinal rows larger than laterals, of irregular shape, pointing posterodorsally; laterals pointing posteroventrally; laterals and ventrals of similar size; ventrals slightly irregular; a row of enlarged scales between supralabials and orbit, bordered by one or two smaller scale rows; gular scales smaller than ventrals, weakly keeled; gular pouch present in males; antehumeral fold/pit weakly developed; nuchal crest composed of 11-15 lanceolate spines; dorsal crest a denticulate ridge; enlarged conical lamellae under the leading edge of third toe; 27-30 lamellae under fourth toe; hind limb length 70-75 % of svl; tail length 238-265% of svl, slightly compressed at the base (Harikrishnan and Vasudevan, 2013). The species within Notacalotes gen. ...
... This genus Notacalotes gen. nov. is alternatively defined and separated from all other agamidae as follows: relatively long head (hw: hl = 0.59); 56-67 longitudinal scale rows around midbody; dorsals and laterals smooth, sometimes weakly keeled near the sacral region; ventrals strongly keeled; dorsals of 4-7 paravertebral longitudinal rows larger than laterals, of irregular shape, pointing posterodorsally; laterals pointing posteroventrally; laterals and ventrals of similar size; ventrals slightly irregular; a row of enlarged scales between supralabials and orbit, bordered by one or two smaller scale rows; gular scales smaller than ventrals, weakly keeled; gular pouch present in males; antehumeral fold/pit weakly developed; nuchal crest composed of 11-15 lanceolate spines; dorsal crest a denticulate ridge; enlarged conical lamellae under the leading edge of third toe; 27-30 lamellae under fourth toe; hind limb length 70-75 % of svl; tail length 238-265% of svl, slightly compressed at the base (Harikrishnan and Vasudevan, 2013). The species within Notacalotes gen. ...
A logical new taxonomy for the Asian subfamily Draconinae based on obvious phylogenetic relationships and morphology of species (Squamata: Sauria: Agamidae: Draconinae)
Article
  • Jul 2014
A number of recent molecular studies have highlighted divergences between south-east Asian agamid lizards within the Draconinae previously thought to be sufficiently close as to be placed in the same genera. Consolidating recently published studies and further investigations into relevant taxa, incorporating available molecular, morphological and geological evidence some genera as presently recognized are re-arranged to better reflect the relationships of species. For some of these taxa, names are available and as a result of this review they are formally resurrected from synonymy. Where names for taxa don’t exist, the species groups are formally named and defined herein according to the Zoological Code (Ride et al. 1999). Well established genera that are in effect split up include the following: Gonocephalus, Kaup, 1825 is divided into three genera. The nominate genus is also split into five subgenera, all named for the first time except for (Dilophyrus Gray, 1845) and the nominate subgenus. Japalura Gray, 1853 is divided into three genera, one of which is named for the first time. Two genera are further divided into two subgenera. Calotes Daudin, 1802 is divided into three genera, two also divided into subgenera. The taxon originally described as “Calotes andamanensis Boulenger, 1891” is also placed in a new monotypic genus as it also clearly sits ouside the two genera in which it has been recently placed (Calotes and Pseudocalotes Kaup, 1827). Ceratophora Gray, 1835, is divided in three, with the two most divergent taxa, C. aspera Günther, 1864 and C. karu Pethiyagoda and Manamendra-Arachchi, 1998 each placed in a new monotypic genus. The remaining Ceratophora are split into two obvious subgenera. Bronchocela Kaup, 1827 is divided into two subgenera. The taxon currently known as Bronchocela cristatella Kuhl, 1820 from the island of Halmahera, Indonesia is formally described as a new species. Other potential new species are identified. Phoxophrys Hubrecht, 1881 is divided into three subgenera, one formally named for the first time. Aphaniotis Peters, 1864 is subdivided into two subgenera as is ptyctolaemus Peters, 1864. Salea Gray, 1845 is divided into two subgenera for which a names are already available. Draco Linnaeus, 1758 is herein conservatively divided nine ways into subgenera of which five are formally named for the first time. It is likely that other taxonomists may treat these divisions as full genera. Furthermore, Draconinae taxonomy and nomenclature at the level between subfamily and genus is tidied up. The result is the formal erection of ten tribes and the addition six subtribes formally defined and named. As a result of this scientific reorganisation of the subfamily, this paper presents a list of all recognized Draconinae species in correct tribes, subtribes, genera and subgenera. Keywords: Taxonomy; nomenclature; genera; Gonocephalus; Japalura; Cophotis; Calotes; Ceratophora; Diploderma; Aphaniotis; Phoxophrys; Bronchocela; Lophocalotes; Dendragama; Otocryptis; Ptyctolaemus; Mictopholis; Salea; Sitana; Otocryptis; Pseudocalotes; Paracalotes; Draco; Dilophyrus; Oriotiaris; Pelturagonia; Lyriocephalus; Coryphophylax; Ptyctolaemus; Mantheyus; Acanthosaura; new genera; Daraninagama; Doongagama; Maxhoseragama; Crottyagama; Skrijelus; Notacalotes; Pethiyagodaus; Manamendraarachchius; subgenera; Dracontoides; Rhacodracon; Pterosaurus; new subgenera; Honlamagama; Mantheysaurus; Denzeragama; Eksteinagama; Jamesschulteus; Rubercalotes; Ghatscalotes; Laccadivecalotes; Ceyloncalotes; Tamilnaducalotes; Freudcalotes; Khasicalotes; Amboncalotes; Ferebronchocela; Olorenshawagama; Proboscisagama; Mindatagama; Macguiredraco; Philippinedraco; Engannodraco; Somniadraco; Spottydraco; new species; Harradineus. new tribes; Dracoiini; Maxhoseragamiini; Crottyagamiini; Daraninagamaiini; Pethiyagodaiini; Japaluraiini; Lophocalotesiini; Phoxophryiini; Mantheyiini; Dendragamaiini; new subtribes; Maxhoseragamiina; Sitanaiina; Acanthosauriina; Saleaiina; Pethiyagodaiina; Doongagamaiina.
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... Mahony (2010) transferred Calotes kingdonwardi Smith, 1935, Japalura kaulbacki Smith, 1937, Mictopholis austeniana (Annandale, 1908 and Salea kakhienensis (Anderson, 1879) to Pseudocalotes and considered J. kaulbacki to be a junior subjective synonym of P. kingdonwardi. Harikrishnan & Vasudevan (2013) rediscovered Calotes andamanensis Boulenger, 1891, and transferred it to Pseudocalotes. Following these changes, the genus currently contains fifteen species. ...
... This scale has not been mentioned in most recent descriptions of congeners and existing photographs of Pseudocalotes have insufficient resolution to score this character. The holotype and one new specimen of P. andamanensis illustrated by Harikrishnan & Vasudevan (2013, their figure 2A and 2B) clearly show heavily fractured scales in the parietal region and no differentiated interparietal in this species. ...
... Unfortunately, color of the buccal epithelium has not been reported for most congeners. In P. larutensis the tongue is red, contrasting with a black throat (Hallermann & McGuire 2001), and it is bright yellow in P. andamanensis (Harikrishnan & Vasudevan 2013). A photo of P. austeniana posted on the Reptile Database (Uetz 2014) reveals that this species has an orange tongue. ...
Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia
Article
Full-text available
  • Jul 2014
We describe three new species of Pseudocalotes from the Bukit Barisan Range of southern Sumatra, Indonesia. Pseudocalotes cybelidermus, P. guttalineatus, and P. rhammanotus differ from most congeners in having serrate dorsal crests that extend to the base of the tail and a dorsolateral series of enlarged heavily keeled scales. In these new species, subdigital lamellae of Toe III have prominent preaxial keels and lack or have greatly reduced postaxial keels. In contrast, P. rhammanotus resembles P. tympanistriga by having bicarinate subdigital lamellae at the base of Toe III. Like most congeners, these new species appear to be restricted to humid forests above 1000 m. We report several new morphological characters for Pseudocalotes and discuss their diagnostic value. Future systematic studies of this genus should assess presence/absence of interparietals, four different kinds of modified scales on the neck, a dorsolateral series of heavily keeled scales, and unicarinate lamellae under the distal phalanges of most fingers and toes. Our comparisons among congeners demonstrate the diagnostic value of width of the gap between the nuchal and dorsal crests and frequency data for contact between the nasal and supralabials and between the postmentals and infralabials. Finally, we discuss variation in morphology of subdigital lamellae at the base of Toe III and describe new conditions intermediate between the serrate fringe of most Indochinese species and the bicarinate lamellae of the P. tympanistriga.
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... Morphological characters were obtained from the newly collected specimens and many key references [1][2][3][4][5][6][7][22][23][24][25][26][27][28][29][30][31][32][33][34][35][36][37]. Measurements and scale counts were performed following Zhao et al., 1999, Harvey et al., 2014, Grismer et al., 2016 and were developed using digital calipers to the nearest 0.1 mm. ...
A New Species of the Genus Pseudocalotes (Squamata: Agamidae) from Southwest Yunnan, China
Article
Full-text available
  • Mar 2024
In this study, a new species of the genus Pseudocalotes is described from Yingjiang County, Dehong Dai and Jingpo Autonomous Prefecture, Yunnan Province, China, based on four female specimens. It can be distinguished from its congeners by the following combination of characters: (1) interoculabials 3 or 4; (2) canthals 5–7; (3) cicrcumorbitals 8–11; (4) 1 scale between rostral and nasal; (5) interparietal 1; (6) superciliaries 4–6; (7) supralabials 6–7, the 1st in contact with the nasal; (8) infralabials 6–8; (9) transverse gular fold and antehumeral fold present; (10) 2–3 enlarged scales between eye and ear; (11) nuchal crest single, consists of 3–5 erected spines; (12) dorsal crest row single, discontinuous and low, located between two keeled, parallel and enlarged scale rows; (13) enlarged postrictals absent; (14) scales around midbody 53–62, dorsal body scales heterogenous in size and shape; (15) midventrals smaller than dorsals; (16) subdigital scales on the 4th finger 20–26, and on the 4th toe 24–29; (17) dorsal background coloration light taupe with four irregular brown patches along the middle of dorsal; (18) inner lips wathet, tongue aurantiacus, throat bluish black. The population from Yingjiang County was nested within a highly supported lineage, formed a sister taxon with P. kakhienensis (SH 97/UFB 100) and according to the p‑distance, the new species differed from its congeners by 14.5% to 35.2% for NADH dehydrogenase subunit 2 (ND2) and 15.5% to 25.0% for NADH dehydrogenase subunit 4 (ND4).
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... For example, no reliable information on locomotion types was available regarding three species of Pseudocalotes sampled in this study. However, Hallermann and McQuire (2001) and Harikrishnan and Vasudevan (2013) both reported slow-moving locomotion of congeners, so we scored the three species of Pseudocalotes sampled in this study as slow-moving. ...
Phylogenetic position of a bizarre lizard Harpesaurus implies the co-evolution between arboreality, locomotion, and reproductive mode in Draconinae (Squamata: Agamidae)
Article
  • Aug 2020
Viviparous reproduction has evolved independently more than 100 times in the evolutionary history of Squamata (lizards and snakes). Adaptation to cold climates is the dominant hypothesis explaining shifts to viviparity, but viviparous species are also present in the tropical lowlands, implying the presence of other factors that may also promote the evolution of viviparity. For example, the tropical Asian/Oceanian subfamily, Draconinae, includes two viviparous genera (Cophotis and Harpesaurus). However, one of them, Harpesaurus, is extremely rare, making it difficult to study aspects of their ecology or evolution. We managed to collect a H. borneensis, and this provided an opportunity to address the evolution of viviparity in draconine lizards. Based on a new molecular phylogenetic hypothesis, including all viviparous groups in Draconinae, we infer that viviparity has evolved twice within the subfamily. Ancestral state reconstruction analyses indicated that shifts to viviparity were preceded by the evolution of arboreality and slow-moving locomotion. Our analysis strongly suggests evolutionary correlations between habitats, locomotion, and reproductive modes. Taken together, these results suggest that the use of arboreal habitats and slow locomotion facilitated the evolution of viviparity in these tropical/subtropical lizards. Arboreal, slow-moving species are considered to rely more on crypsis in predator avoidance rather than fleeing. Therefore, we propose that such species are relatively insusceptible to the concomitant cost of viviparity: reduced locomotive ability and increased risk of predation during pregnancy. We also describe and discuss the unique behaviours of H. borneensis.
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... Since Smith (1935), there have been several changes to the classification of the Indian and Indo-Chinese agamids. Now all these agamids belong to subfamily Draconinae; Smith's group I: C. cristatella, C. jubatus, C. smaragdinus are all under the genus Bronchocela (Moody, 1980); group II: C. microlepis, C. floweri are now under the genus Pseudocalotes (Moody, 1980) while C. fruhstorferi was moved to Acanthosaura (Denzer et al. 1997); group III: C. versicolor, C. maria, C. jerdoni, C. emma, C. mystaceus, C. nemoricola, C. grandisquamis and C. calotes remain in the genus Calotes while C. kakhienensis was moved to the genus Pseudocalotes (Mahony, 2010); group IV: C. liocephalus, C. liolepis, C. nigrilabris and C. ceylonensis remain in the genus Calotes while C. kingdom-wardi was moved to the genus Pseudocalotes (Mahony, 2010) and C. andamanensis was moved to the genus Pseudocalotes (Harikrishnan & Vasudevan, 2013) but with a cautionary note that this species has a weak ante-humeral fold which is otherwise present in genera such as Calotes, Psammophilus, Gonocephalus, Japalura and Acanthosaura; group V: C. ellioti and C. rouxii have been moved to the new genus Monilesaurus. This study has added more stability to the systematics of Draconinae particularly Calotes and other endemic agamids from the Western Ghats using comprehensive data on their morphology and genetics. ...
A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species
Article
Full-text available
  • Jul 2018
Lizards of the genus Calotes are geographically restricted to South Asia, Indo-China and parts of Southeast Asia. The greatest diversity of the genus is from the biodiversity hotspots in South Asia: Western Ghats (Peninsular India), Sri Lanka and Indo-Burma. Here, we present a systematic revision of members of the genus Calotes from Peninsular India using a combination of molecular phylogeny, geographical distribution and morphological characters. We show that Calotes from the Western Ghats is paraphyletic and consists of three major clades, one of which is widely distributed in South and Southeast (SE) Asia, while the others are restricted to Peninsular India. The Peninsular Indian clade is composed of two sister clades: Psammophilus, with a wider distribution and a second clade, composed of two extant species, Calotes rouxii and Calotes ellioti and two new species, all restricted to the Western Ghats region. Based on morphological differences, we retain the generic status of Psammophilus and assign its sister clade to a new genus Monilesaurus gen. nov. and transfer the following species, C. rouxii and C. ellioti, to this new genus. We also provide diagnoses and descriptions for two new species recognized within Monilesaurus gen. nov. In addition, Calotes aurantolabium from the Western Ghats was observed to be deeply divergent and to share a sister-relationship with the clade composed of Calotes, Monilesaurus gen. nov., and Psammophilus. Based on its phylogenetic position and morphological attributes, we assign this species to a new genus Microauris gen. nov. These new discoveries highlight the evolutionary significance of the Western Ghats in housing novel lizard diversity.
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... Singapore has several low hills, with Bukit Timah being the highest at only 164 m. Although most species of Pseudocalotes occur above 800 m, at least one insular species, P. andamanensis, occurs near sea level (Harikrishnan and Vasudevan 2013). ...
Endemic Dragons of Sumatra's Volcanoes: New Species of Dendragama (Squamata: Agamidae) and Status of Salea rosaceum Thominot
Article
Full-text available
  • Dec 2017
  • HERPETOL MONOGR
We discovered new populations of Dendragama at the northern and southern ends of Sumatra. High genetic distances and concordance of multiple, apparently independent diagnostic characters support our descriptions of these two populations as new species. We define new characters of the sublabial, tympanic, dorsal crest, and dorsolateral crest scales. The three species of Dendragama undergo remarkable color change in response to time of day and stress. Females lay 2–4 ovoid eggs, reach sexual maturity at about 60 mm snout–vent length, and likely produce multiple clutches each year. We remove Salea rosaceum Thominot from the synonymy of Dendragama boulengeri and argue that the unique holotype of S. rosaceum is a specimen of Pseudocalotes tympanistriga with incorrect locality information.
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... for Bronchocela,McGuire et al. (2007) for Draco,Denzer & Manthey (2009) for Gonocephalus,Mahony (2009) for Japalura,Zug et al. (2006),Krishnan (2008) andHallermann (2000) for Calotes,Harikrishnan & Vasudevan (2013) for Pseudocalotes,Pethiyagoda & Manamendra-Arachchi (1998) for Ceratophora andSchulte et al. (2004) for Ptyctolaemus. Several genera such as Harpesaurus, Thaumatorhynchus, and Psammophilus are not treated at all. ...
A critical review of Hoser’s writings on Draconinae, Amphibolurinae, Laudakia and Uromastycinae (Squamata: Agamidae)
Article
Full-text available
  • Mar 2016
We analyzed four papers on agamid lizards by self-proclaimed Australian herpetologist Raymond Hoser with respect to the presentation of diagnostic characters as well as their taxonomic and nomenclatural merits. In most cases the taxonomic concepts were lifted from earlier phylogenetic publications and the diagnoses were copied from other authors. Copied text in Hoser’s diagnostic section within the analyzed papers amounts to a staggering 83% for Draconinae, 82% for Amphibolurinae, 77% for Laudakia and 78% for Uromastycinae, respectively. We found a number of plagiarized paragraphs, sometimes half a page long. Hoser hardly ever makes any effort to attribute statements to the original author and in some cases he even omitted to cite the relevant source. With respect to nomenclature, we found that Hoser proposed names that were preoccupied or unavailable, that a nomen oblitum was resurrected incorrectly, nomina nuda were produced, a type locality was restricted incorrectly and a questionable holotype was designated for a new species. With respect to taxonomy, we found examples of wrong diagnoses, falsely attributed species, omission of taxa and a lack of understanding or misinterpretation of previously published taxonomic studies on agamid lizards. Furthermore relevant literature on taxonomy and nomenclature has been overlooked or disregarded.
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Dragons in the mist: Three new species of Pseudocalotes Fitzinger (Squamata: Agamidae) from the sky island archipelago of Peninsular Malaysia
Article
  • Jul 2016
An integrative taxonomic analysis is used to delimit and describe three new species of Pseudocalotoes from the sky island archipelago of the Banjaran (=mountain range) Titiwangsa of Peninsular Malaysia. Pseudocalotes drogon sp. nov., from Fraser’s Hill, Pahang is basal to the sister species P. larutensis from Bukit Larut, Perak in the Banjaran Bintang and the new species P. rhaegal sp. nov. from Cameron Highlands, Pahang. Pseudocalotes drogon sp. nov. is differentiated from all other species of Psuedocalotes by having the combination of a flat rostrum; seven postrostrals; an interparietal; 11 cir- cumorbitals; five canthals; 7–10 superciliaries; one scale between the rostral and nasal; nine supralabials; eight infralabi- als; 10 postnasal-suborbital scales; four postmentals; five or six sublabials; five or six chinshields; 47 smooth, wide, gular scales; weak transverse gular and antehumeral folds; two enlarged scales between the ear and eye; enlarged upper and low- er posttemporals; a single enlarged supratympanic; no enlarged postrictals; three large scales bordering the dorsal margin of the ear opening; large pretympanic scales; eight scales in the nuchal crest not separated by a gap; enlarged vertebral scales extending to the tip of the tail; keeled and non-plate-like scales on flanks; 51 midbody scales; midventrals smaller than dorsals; 19 subdigital lamellae on the fourth finger; 23 subdigital lamellae on the fourth toe; preaxial scales on third toe enlarged and spinose; subdigital lamellae not unicarinate; HW/HL 0.52; HL/SVL 0.31; no elbow or knee patches; and a male dewlap color of lime-green bearing a central yellow spot. Pseudocalotes rhaegal sp. nov. is differentiated from all other Psuedocalotes by having the combination of a convex rostrum; 6–8 postrostrals; an interparietal; nine or 10 circu- morbitals; five canthals; 7–10 superciliaries; one or two scales between the rostral and nasal scales; eight or nine suprala- bials; seven or eight infralabials; 11 or 12 postnasal-suborbital scales; four postmentals; four or five chinshields; 40–45 smooth, wide, gular scales; no transverse gular fold; a weak antehumeral fold; three or four enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; no large scales bordering the upper margin of the ear opening or in the pretympanic region; 6–8 enlarged nuchal crest scales not separated by a gap; enlarged vertebral scales extending to the base of the tail; weakly keeled, non-plate-like scales on the flanks; 52– 58 midbody scales; midventrals smaller than dorsals; 19–21 subdigital lamellae on the fourth finger; 22–26 subdigital la- mellae on the fourth toe; preaxial scales on the third enlarged and rounded; subdigital lamellae not unicarinate; HW/HL 0.50–0.54; HL/SVL 0.28–0.30; no elbow or knee patches; and female dewlap color yellow bearing a purple base. The analyses also indicated that the new species, P. viserion sp. nov. from Genting Highlands, Pahang in the southern section of the Banjaran Titiwangsa is the sister species of P. flavigula from Cameron Highlands 121 km to the north and can be separated from all other species of Psuedocalotes by having the combination of three postrostrals; 10 circumorbitals; four or five canthals; 5–7 superciliaries; rostral and nasals in contact; supralabials contacting the nasal; six or seven suprala- bials; six or seven infralabials; two or three postmentals; 47 or 48 smooth, flat, gular scales; three chinshields; weak trans- verse gular and antehumeral folds; two enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; 7–9 nuchal crest scales lacking gaps and not extending beyond midbody; weakly keeled and plate-like scales on the flanks; 35–38 midbody scales; ventrals smaller than dorsals; 22 or 23 subdigital lamellae on the fourth finger; 26 or 27 subdigital lamellae on the fourth toe; preaxial scales on the third toe not modified; subdigital scales not unicarinate; HW/HL 0.62; no white marking below the eye; dewlap in males yel- low; and no elbow or knee patches. Pseudocalotes rhaegal sp. nov. most likely occurs in syntopy with P. flavigula in Ta- nah Rata at Cameron Highlands and its discovery adds to a growing body of literature detailing the recent descriptions of several new, upland, closely related, sympatric species in Peninsular Malaysia. Another new population referred to here as Pseudocalotes sp. nov. from the Hala-Bala Wildlife Sanctuary, Betong District, Yala Province, Thailand is discussed. The discovery and description of these three new Pseudocalotes from the upland regions of Peninsular Malaysia continues to underscore the remarkably high herpetological diversity and ecological complexity in this sky island archipelago that is still underestimated, unappreciated, and unprotected.
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A review of the genus Pseudocalotes (Squamata: Agamidae), with description of a new species from West Malaysia
Article
Full-text available
  • May 2000
A new species of the genus Pseudocalotes (P. dringi n. sp.) from the mountain forest of Western Malaysia is described. The holotype of the new species was formerly regarded as a syntype of P. floweri. A review of all species of the genus is given including a differential diagnosis and a key to the species. Diagnostic characters for the genera Bronchocela, Calotes, Dendragama were established. Paracalotes poilani and Pseudocophotis sumatrana, showing all diagnostic characters of the genus Pseudocalotes, are included in that genus. Members of Pseudocalotes of the Indochinese region can be differentiated from members inhabiting the Sunda region by the shape of the scales under the third toe.
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Maps of Pleistocene sea levels in Southeast Asia: Shorelines, river systems and time durations
Article
Full-text available
  • Sep 2000
  • J BIOGEOGR
AimGlaciation and deglaciation and the accompanying lowering and rising of sea levels during the late Pleistocene are known to have greatly affected land mass configurations in Southeast Asia. The objective of this report is to provide a series of maps that estimate the areas of exposed land in the Indo-Australian region during periods of the Pleistocene when sea levels were below present day levels.LocationThe maps presented here cover tropical Southeast Asia and Austral-Asia. The east–west coverage extends 8000 km from Australia to Sri Lanka. The north–south coverage extends 5000 km from Taiwan to Australia.Methods Present-day bathymetric depth contours were used to estimate past shore lines and the locations of the major drowned river systems of the Sunda and Sahul shelves. The timing of sea level changes associated with glaciation over the past 250,000 years was taken from multiple sources that, in some cases, account for tectonic uplift and subsidence during the period in question.ResultsThis report provides a series of maps that estimate the areas of exposed land in the Indo-Australian region during periods of 17,000, 150,000 and 250,000 years before present. The ancient shorelines are based on present day depth contours of 10, 20, 30, 40, 50, 75, 100 and 120 m. On the maps depicting shorelines at 75, 100 and 120 m below present levels the major Pleistocene river systems of the Sunda and Sahul shelves are depicted. Estimates of the number of major sea level fluctuation events and the duration of time that sea levels were at or below the illustrated level are provided.Main conclusionsPrevious reconstructions of sea-level change during the Pleistocene have emphasized the maximum lows. The perspective provided here emphasizes that sea levels were at their maximum lows for relatively short periods of time but were at or below intermediate levels (e.g. at or below 40 m below present-day levels) for more than half of each of the time periods considered.
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New Species of Calotes (Reptilia: Squamata: Agamidae) from the Southern Western Ghats, India
Article
  • Sep 2009
A new species is described from the southern Western Ghats of India. The new species was wrongly identified as Calotes andamanensis and is distinguished from it in lacking the antehumeral pit and possessing acutely keeled scales. It is distinguished from all species of the Calotes versicolor group of Smith in having posteroventral orientation of dorsal body scales and is distinguished from all species of the Calotes liocephalus group of Smith in lacking the antehumeral pit. A redescription of C. andamanensis is provided based on the holotype. Calotes andamanensis is a valid species with type locality in the Andaman and Nicobar Islands, India.
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Phylogeny of Cerberus (Serpentes: Homalopsinae) and phylogeography of Cerberus rynchops: diversification of a coastal marine snake in Southeast Asia
Article
  • Aug 2004
  • J BIOGEOGR
Aim The biogeography of Southeast Asia has been greatly affected by plate tectonic events over the last 10 Myr and changing sea levels during the Quaternary. We investigated how these events may have influenced the evolution of Cerberus Cuvier, a marine coastal snake belonging to the Homalopsinae (Oriental‐Australian Rear‐fanged Water Snakes). This study is an expansion of a previous study on the biogeography and systematics of Cerberus . Location We obtained species from localities across the range of the widely distributed Cerberus : India, Sri Lanka, the Andaman islands, Myanmar, the Philippines, Borneo, Suluwesi, Sumatra, Vietnam, Thailand, Singapore and Australia. Methods We analysed mtDNA sequences (12S, ND3, ATPase, 2338 nucleotide characters) from 21 localities. The sample consisted of 65 Cerberus rynchops (Schneider), three Cerberus australis (Gray) and four Cerberus microlepis Boulenger. One Homalopsis buccata (Linnaeus), one Bitia hydroides Gray, one Enhydris enhydris (Schneider), and two Enhydris plumbea (Boie) were used as outgroups. Results We produced phylogenetic trees based on parsimony, maximum likelihood and Bayesian analysis. We did not find unambiguous support for the monophly of Cerberus . Cerberus austalis , H. buccata and all other Cerberus populations formed a three‐way basal polytomy under parsimony and C. australis formed the sister group to a clade consisting of H. buccata and all other Cerberus in likelihood and Bayesian analysis. The non‐Australian Cerberus were monophyletic and consisted of four primary biogeographical clades: Indian and Mayanmar, Philippines, Greater Sunda Islands and Suluwesi, and the Thai‐Malay peninsula and Gulf of Thailand. The range of genetic divergence between these clades and Australian Cerberus was 0.06–0.12. Genetic divergence among clades to the west of Australia was less pronounced (Thai‐Malay peninsula and Gulf of Thailand = 0.02–0.05; Sunda Islands and Suluwesi = 0.02–0.05; Philippines = 0.02–0.06; India and Myanmar = 0.04–0.06, Philippines = 0.02–0.5). Main conclusions Gyi [ University of Kansas Publications, Museum of Natural History 20 (1970), 47] recognized three species of Cerberus : C. australis (from Australia), C. microlepis (known only from Lake Buhi in the Philippines), and the widely distributed C. rynchops (India to Wallacea). We did not find strong support for the monophyly of the genus. Cerberus australis is highly divergent from all other Cerberus lineages sampled from this region. The geographically widespread C. rynchops is resolved into four biogeographical clades (Indian and Myanmar, Philippines, Greater Sunda Islands and Suluwesi, and the Thai‐Malay Peninsula and Gulf of Thailand). We discuss how the dispersal biology of a salt‐water tolerant, coastal marine taxon and the complex geological history of the region (Tertiary plate tectonic movements and Quaternary sea‐level changes) could produce the observed patterns of diversification.
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Bronchocela cristatella
  • Jan 1820
ZsIC – Zoological survey of India (Kolkata, India). Bronchocela cristatella (Kuhl, 1820)
ZsI 3880-sK03nC1, nan - cowry; Calotes aurantolabium KrIshnAn
  • Mann
MAnn, 2009 – ZsI 3880-sK03nC1, nan - cowry; Calotes aurantolabium KrIshnAn, 2008 – BnhM 1436, (holotype), Kalakkad Mundanthurai Tiger reserve, Tamil nadu, India; Calotes jerdoni (gunTher, 1870) –
Biogeography of the amphibians and reptiles of the Andaman and nicobar Islands, India
  • Jan 1999
  • 43-77
  • I Das
DAs, I. (1999): Biogeography of the amphibians and reptiles of the Andaman and nicobar Islands, India; pp. 43-77. In: oTA, h. (ed.) Tropical island herpetofauna. origin, current diversity and current status. Amsterdam (elsevier).
with a short account of the natives, their customs and habits, and of previous attempts at colonization; Calcutta (office of the superintendent of government printing), pp. 140. hAllerMAnn
  • Jan 1875
  • J HERPETOL
  • 255-265
  • F A De Roepstorff
  • J Hallermann
  • J A Mcguire
De roepsTorFF, F. A. (1875): Vocabulary of dialects spoken in the nicobar and Andaman Isles, with a short account of the natives, their customs and habits, and of previous attempts at colonization; Calcutta (office of the superintendent of government printing), pp. 140. hAllerMAnn, J. (2005): A taxonomic review of the genus Bronchocela (squamata: Agamidae) with the description of a new species from Vietnam.russian Journal of herpetology, Moscow; 12 (3): 167-182. hAllerMAnn, J. & BöhMe, w. (2000): A review of the genus Pseudocalotes (squamata: Agamidae), with description of a new species from west Malaysia.Amphibia-reptilia, leiden; 21: 193-210. hAllerMAnn, J. & MCguIre, J. A. (2001): A new species of Pseudocalotes (squamata: Agamidae) from Bukit larut, west Malaysia.-herpetologica, lawrence; 57 (3): 255-265.
rediscovery of Calotes andamanensis Boulenger (1891) and a reassessment of the type locality
  • Jan 1998
  • 530-535
  • N M Ishwar
  • I Das
IshwAr, n. M. & DAs, I. (1998): rediscovery of Calotes andamanensis Boulenger (1891) and a reassessment of the type locality.-Journal of the Bombay natural history society, Mumbai; 95: 513-514. KrIshnAn, s. (2008): new species of Calotes (reptilia; squamata: Agamidae) from the southern western ghats, India.-Journal of herpetology, houston etc.; 42 (3): 530-535.