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New Species of Calotes (Reptilia: Squamata: Agamidae) from theSouthern Western Ghats, India

SHREYAS KRISHNAN1

Department of Biology, University of Texas at Arlington, Texas 76019, USA

ABSTRACT.—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 Calotesliocephalus 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.

The Western Ghats, a mountain range run-ning parallel to the west coast of the Indianpeninsula, extend approximately 1,500 km al-most continuously from 22u–8uN (Fig. 1). Themountain range is wider in the north and taperstoward the tip of the peninsula and is inter-rupted by a 30-km break at the Palghat Gap at10u339N and the narrower Shencottah Pass(9uN). South of 11uN the mountains receiverainfall from both the southwest and northeastmonsoon systems through nine months of theyear; consequently, the region is also among thehighest in floral and faunal endemicity (Ishwar,2001). The Kalakad Mundanthurai Tiger Re-serve (KMTR, 895 km2, 8u259–8u539N and77u109–77u359E) is located at the southern tipof the Western Ghats, from 50–1,800 m altitude.Rain forests within the reserve extend from 700–1,600 m and have an annual precipitationranging from 750–3,000 mm (Ishwar et al.,2001). The most recent published herpetofaunalwork in KMTR was conducted by the WildlifeInstitute of India. During their surveys, anarboreal agamid was discovered and later

identified by Ishwar and Das (1998) to beCalotes andamanensis Boulenger, 1891. Ishwarand Das (1998) concluded that C. andamanensis

 belongs to the peninsular Indian and Sri Lankanlineage of  Calotes and accordingly emended itsdistribution.

The specific epithet of  C. andamanensis refersto its original collection locality in the AndamanIslands (Fig. 1). The Andaman and NicobarIslands are archipelagos approximately 1,200 kmeast off the southeast coast of India runningnorth-south along a submarine ridge between

Myanmar and Sumatra (6u–14uN and 92u–94uE). The Andaman and Nicobar archipelagosare comprised of 306 islands, islets, and rocks.The northern Andaman Islands are separatedfrom the southern Nicobar Islands by a distanceof 140 km and from the Isthmus of Kra by650 km of Andaman Sea at 10uN. Floral andfaunal distributions suggest an Indo-Myanmaraffinity for the Andamans and oceanic condi-tions for the Nicobars based on their depauper-ate flora and fauna (Das, 1999). The holotype of C. andamanensis (ZMUC 36944) was collected by

F. A. de Roepstorff in 1882 (J. Rasmussen, pers.comm.; Singh, 2003) and described by G. A.Boulenger in 1891. Ishwar and Das (1998)compared specimens from KMTR to a damagedspecimen misidentified as C. andamanensis(BNHM 710) in the collections of the BombayNatural History Society, India, and with thedescription of  C. andamanensis given in Smith(1935); the holotype was not examined. Thispaper assigns the specimens identified as C.andamanensis (from KMTR) by Ishwar and Das(1998) to a new species and provides a rede-

scription of C. andamanensis.

MATERIALS AND METHODS

Museum abbreviations follow Leviton et al.(1985). The following specimens were examined(see Appendix 1): C. andamanensis, identified byIshwar and Das (1998), and the holotype C.andamanensis Boulenger, 1891. Representativesof the genus Calotes that occur in KMTR, southIndia, and Sri Lanka were examined: Calotesliolepis, Calotes ceylonensis, Calotes calotes, Calotes

rouxi, Calotes nemoricola, Calotes nigrilabris, andCalotes versicolor. Calotes desilvai, Calotes ellioti,and Calotes liocephalus were compared based onliterature (Bahir and Maduwage, 2005; Smith,1935) and field examinations of  C. ellioti.

1 Present address: 2633 McKinney Avenue,#130-352,Dallas, Texas 75204, USA; E-mail: shreyas.krishnan@gmail.com

 Journal of Herpetology, Vol. 42, No. 3, pp. 530–535, 2008Copyright 2008 Society for the Study of Amphibians and Reptiles

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Specimens were examined for meristic andmetric morphological characters. Morphometricdata were recorded using digital calipers(0.03 mm precision rounded to the nearest0.01 mm), and mass was recorded using a 30gPesola scale (6 0.3% of load). Meristic charac-ters included number of scales along themidventral torso (between axilla and femur)

and number of scales around the midbody.Labial scales (upper and lower) were recordedon the left and right sides, whereas theremaining characters were recorded on eitherthe right or the left side of the body. Canthals,continuous with superciliary scales, from thelast superciliary scale up to the nasal. Suborbi-tals, from the canthal edge to the level of the lastsuperciliary scale/postcircumorbital. Circumor-

 bitals, from the canthal around the dorsal edgeof the orbit up to the last superciliary scale/postcircumorbital. Number of scale rows be-tween tympanum and orbit, from the first rowof scales abutting the tympanum up to thesuborbital ridge. Subdigital lamellae (first,fourth, and fifth digits of the pes), from the firstlamella at the base of the toe up to the claw.

Snout–vent length (SVL) was recorded afterdrying the specimen of excess alcohol. Headlength (HL), distance from posterior margin of the tympanum to the front surface of the rostral,head width (HW), distance between the angle of the lips, measured dorsally to exclude the jaw

musculature, and head height (Hht), at theangle of the lips from the ventral face of theflattened throat up to the parietal region. Nasalwidth (NW), distance between nasals; nasalheight (NH), snout height at the nasals from thelower surface of the dentary to the canthalsdirectly above. Limb measurements, from axil-la–elbow or groin–knee for the humerus (Hu-merus) and femur (Femur) length, respectively,and antebrachial (Ulna) and shank (Tibia)lengths were measured up to the carpals ortarsals, respectively. Shoulder height (ShHt), at

the level of the humeral tuberosities up to themiddorsal at the level of the vertebral scales,and hip height (HpHT), at the pelvic girdle fromthe venter up to the ilial crests. Foot length (FL),from the tarsals to the claw tip on the longesttoe. Tympanum diameter (TD), distance be-tween anterior margin of tympanum andposterior margin of orbit (TO), and horizontallength of orbit (O) were measured. Tympanumdiameter was taken as the average of thehorizontal and vertical diameters. Mass wasalso recorded after drying the specimen of 

excess alcohol.

SYSTEMATICS

Calotes aurantolabium sp. nov.

Calotes andamanensis Ishwar and Das (1998) part

 Holotype.—Bombay Natural History Society(BNHM) 1436 (Orange-lipped forest lizard) anadult female (Fig. 2 A, B, E, G), collected by N.M. Ishwar in Kalakad Mundanthurai TigerReserve (Fig. 1), Tamil Nadu, India in 1997.

Diagnosis.—Calotes aurantolabium is diagnosed by having an orange streak above each row of supralabials, green body; acutely keeled scalesover body (dorsally and ventrally), head, andthroat; posteroventral orientation of the dorsalscales; antehumeral pit absent; 63 scales aroundmidbody; small tympanum (5.5% HL); toe IIIand IV subequal. Distinguished from C. anda-manensis in having acutely keeled dorsals, all of which are directed posteroventrally; antehum-eral pit absent; acutely keeled ventrals, limb andhead scales; smaller occipital, nuchal, temporalregions. Distinguished from all known speciesof the C. versicolor group in having poster-oventral orientation of dorsal scales (poster-odorsal in C. versicolor group). Distinguishedfrom species of the C. liocephalus group in

FIG. 1. Relief map of southern India showing typelocality, Kalakad Mundanthurai Tiger Reserve, of Calotesaurantolabium at 8uN. Elevation above sea level repre-sented in grayscale (white, approximately 0 m, darkgrays to black . approximately 1,000 m). Inset showsregional perspective of type localities of C. aurantolabium,Kalakad Mundanthurai Tiger Reserve (A); and Calotesandamanensis, Andaman and Nicobar Islands (B).

Map rendered from http://www.itis-molinari.mi.it/Gtopo30.htm.

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lacking antehumeral pit, and in having aproportionately smaller head, ulnar lengthproportionately longer, tibial length proportion-ately shorter (unpubl. data). Distinguished fromC. rouxi and C. elliotti in lacking antehumeralfolds and spines; toe-IV longer than III in C.rouxi and C. elliotti.

Description of Holotype.—Adult female; SVL67.88 mm; mass 6.25 g; supralabials 10/10, infra-labials 10/11; circumorbital 15; canthals 13;suborbitals 22; supraorbital rows 6; interorbitals4; palpebrals 22; postmentals 2; tympanumless than one-third eye diameter; tympanum

diameter 5.5% head length or one-fifth distance  between tympanum and orbit; 8 scale rows  between tympanum and orbit, uniform size;dorsal head scales with acute, medial keel; allhead scales uniform in size; labials smooth; threescales surround rostral; nasal undivided;canthals continuous with superciliary scales;antehumeral pit or fold absent; chin and throatscales keeled; throat scales smaller than ventrals;parietal region flat; gular pouch poorly devel-oped; nuchal crest poorly developed, 13 scales;

  body crest poorly developed, not continuouswith nuchal crest; body laterally compressed;

FIG. 2. Right lateral and dorsal head, lateral body, and throat-chin views of  Calotes aurantolabium, female

holotype BNHM 1436 (A, B, E, G) and Calotes andamanensis, male holotype ZMUC 36944 (C, D, F,H), respectively.

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scales around midbody 63; dorsal body scalesuniform, acutely keeled, first row directedposteriorly, remainder posteroventrally; ventralscales acutely keeled as large as dorsals; sub-digital lamellae: toe-I 7, IV 25, V 18; toe-IV and IIIsubequal (IV< III . V . II . I), finger-III equals

IV; limb scales keeled.Variation.—Two females were reported by

Ishwar and Das (1998): SVL 76–79 mm; tail 127–147 mm; mass 7–10 g; supralabials 10 and 10;infralabials 10 and 10. Four eggs were produced

 by both specimens (Ishwar and Das, 1998).Color in Life.—Light green dorsally, suprala-

  bials bluish, orange stripe between supralabialsand orbital, orange stripe on thighs, black spotson trailing edges (Ishwar and Das, 1998).

Color in Preserved Condition.—Orange streakover upper lip, extending over cheek and onto

the shoulder; olive laterally; ventral scalestending to pale brown and white (presentexamination). Supralabials bluish; orange streak

 between supralabial and orbital region; dorsalslight green; orange stripe on thigh; black spotson trailing edges (Ishwar and Das, 1998).

Etymology.—The specific epithet is derivedfrom the Neo-Latin aurantium, meaning golden,and the Latin labium, meaning lip, in referenceto the orange upper lip.

Distribution and Habitat.—The habitat at thetype locality is tropical evergreen rain forest.Sympatric agamids include C. elliotti, C. rouxi,

Calotes grandisquamis, C. nemoricola, Draco dussu-mieri, Otocryptis beddomii, Psammophilus blanfor-danus, and Psammophilus dorsalis (Ishwar et al.,2001). Calotes calotes and C. versicolor occur atlower elevations in drier habitat (Ishwar et al.,2001).

Calotes andamanensis Boulenger (1891)

 Holotype.—Zoological Museum, KøbenhavnsUniversitet, Copenhagen, Denmark (ZMUC)36944 (Fig. 2 C, D, F, H), adult male fromAndaman Islands, India (Fig. 1); collected by

Frederick Adolph de Roepstorff on 7 January1882.

Diagnosis.—Calotes andamanensis is diagnosed by having enlarged keeled scales on posteriorsurface of thigh, dorsal head scales obtuselykeeled, smooth dorsal body scales (upper sixrows directed posterodorsally, remainder pos-teroventrally), and lacking body crest, ante-humeral pit present; nuchal crest well devel-oped; 67 scales around midbody; tympanum(11% HL); toe III shorter than IV. Distinguishedfrom C. aurantolabium in having smooth dorsals,

dorsal body scales unequal, upper six scalerows larger, remainder equal in size to ventralscales; three enlarged scales on caudal thigh;dorsal head scales obtusely keeled; parietalridge raised; enlarged scale between nuchal

crest and tympanum; antehumeral pit present;toe-IV longer than III; stretched hind limbreaches eye. Distinguished from C. versicolorand C. liocephalus groups and C. rouxi and C.elliotti in presence of enlarged keeled scales oncaudal surface of thigh. Distinguished from C.

versicolor group in scale orientation; distin-guished from C. versicolor in having an ante-humeral pit; distinguished from C. nemoricolaand C. grandisquamis in having equal size dorsaland ventral scales, toe-IV longer than III, scalesaround midbody 67 (36–43 and 27–35, respec-tively); distinguished from C. calotes in lackingflattened spines in the nuchal region. Distin-guished from C. elliotti and C. rouxi in having anantehumeral pit (instead of folds) and in lackingspines in the nuchal region. Distinguished fromC. liocephalus group (Calotes ceylonensis, C.

desilvai, C. liolepis, C. nigrilabris) in lackingspines in the nuchal region; distinguished fromC. liocephalus by midbody scale count and bodycrest scales.

Description of Holotype.—Adult male; SVL84.1 mm; mass 12.5 g; supralabials 10/10; infra-labials 10/13; circumorbitals 12; canthals 14/13;suborbitals 16/17; supraorbital rows 5; inter-orbitals 4; palpebrals 27; postmentals 2; tympa-num as large as eye, diameter 11% head lengthor half the distance between tympanum andorbit; cluster of 5 enlarged scales betweentympanum and orbit; 7–8 scale rows between

tympanum and orbit; conical/obtusely keeleddorsal head scales; antehumeral pit present;chin and throat scales smooth; throat scalessmaller than ventrals; body laterally com-pressed; nuchal crest well developed, 15 tallcompressed triangular scales; body crest poorlydeveloped, not continuous with nuchal crest;scales around midbody 67; dorsal body scalesunequal, not keeled, upper six scale rows largerand directed posteriorly and posterodorsally,remainder equal in size to ventral scales anddirected posteroventrally; ventral scales strong-

ly keeled; third and fourth fingers equal;subdigital lamellae: finger-III 22, IV 23, toe-I 9,IV 28, V 20; toe-IV (15.9 mm) longer than III(13 mm) (IV . III . V . II . I); three enlargedscales on caudal thigh; stretched hind limbreaches eye (including toe); tail weakly keeleddorsally, tail strongly keeled ventrally. Morpho-metric comparisons between C. andamanensisand C. aurantolabium are shown in Table 1.

Color in Preservation.—Off white (presentexamination). Top of head yellow, greenishwhite ventrally (Smith, 1935); dorsal body colorgreen with white spots, tail with blackish annuli(Boulenger, 1891).

Remarks.—The holotype was collected byFrederick Adolph de Roepstorff who wasOfficer in Charge of the British camp at

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Nancowry harbor, Nicobar Islands from Janu-ary 1882 onward. The islands of Trinket,

Kamorta and Katchal are a short distance fromthe harbor and may have been visited byRoepstorff. In addition, Roepstorff is known tohave visited Great Nicobar Island. His fieldnotes were not available for examination,though he is not known to have visited theAndaman Islands (M. Chandi, pers. comm.).Calotes andamanensis was presented to theCopenhagen collections upon his return on 1

  June 1887 (M. Andersen, pers. comm.). Themidbody scale count of Boulenger (1891) doesnot match that of Smith (1935) or my observa-

tions. An additional specimen BNHM 710 wasin very poor condition and not diagnosable. Aseparate publication on the systematics of Calotes andamanensis and related new speciesfrom the Andaman and Nicobar Islands is inpreparation.

 Acknowledgments.—Fieldwork and museumvisits were supported by grants from the Societyfor the Study of Amphibians and Reptiles,Explorers Club, Phi Sigma and Department of Biology and College of Science of the Universityof Texas at Arlington, the Madras CrocodileBank Trust (MCBT), and personal funds frommy father. I specially thank V. Giri for bringingthe case of Calotes aurantolabium to my attention,and the curator of the BNHM (N. Chaturvedi)

and their staff for allowing use of their facilities.The curators and collections managers of thecollections at the Field Museum of NaturalHistory (H. K. Voris, A. Resetar), SmithsonianInstitution (K. de. Quieroz and R. Wilson),University of Texas at Arlington (J. A. Camp-

  bell, C. J. Franklin, P. C. Ustach), ZoologicalMuseum, Københavns Universitet, Copenha-gen, Denmark (late J. Rasmussen, M. Andersen),Zoological Survey of India-Kolkata (J. R. B.Alfred, N. C. Gayen, G. Dasgupta) are thankedfor permitting use of their collections. Saw.

  John, Saw. Glan, Saw. Agu, Montu, Saw.Nelson, M. Chandi, and Athi Paungwey in theAndaman and Nicobar Islands are thanked forhosting me, assisting with related collections,specimens, and field trips. P. D. Venugopal,provided some ecological data on Calotes aur-

antolabium. The MCBT staff and H. V. Andrews,for use of their facilities. J. A. Campbell, E. N.Smith, and G. Zug, for critically reading earlierdrafts of the manuscript. I. Das, T. M. Doan, andan anonymous reviewer are thanked for com-ments to improve the final manuscript.

LITERATURE CITED

BAHIR, M. M., AND K . P . MADUWAGE. 2005. Calotesdesilvai, a new species of agamid lizard fromMorningside forest, Sri Lanka. The Raffles Bulletinof Zoology 12(Suppl.): 381–392.

BOULENGER, G. A. 1891. On new or little known Indianand Malayan reptiles and batrachians. Annals andMagazine of Natural History 68:288–292.

DAS, I. 1999. Biogeography of the amphibians andreptiles of the Andaman and Nicobar Islands,India. In H. Ota (ed.), Tropical Island Herpeto-fauna: Origin, Current Diversity, and Conserva-tion, pp. 43–77. Elsevier Science B.V., Amsterdam,The Netherlands.

ISHWAR, N. M. 2001. Reptilian Species Distribution inResponse to Habitat Fragmentation and Microhab-itats in the Rainforests of Southern Western Ghats,India. Unpubl. Ph.D. diss., Forest Research Insti-tute (Deemed University), Uttaranchal, India.

ISHWAR, N. M., AND I. DAS. 1998. Rediscovery of Calotesandamanensis Boulenger (1891) and a reassessmentof the type locality. Journal of the Bombay NaturalHistory Society 95:513–514.

ISHWAR, N . M . , R . CHELLAM, AND A. KUMAR. 2001.Distribution of forest floor reptiles in the rainforestof Kalakad–Mundanthurai Tiger Reserve, SouthIndia; Special Section: Kalakad–MundanthuraiTiger Reserve. Current Science 80:413–418.

LEVITON, A. E., R. H. GIBBS JR. , E. HEAL, AND C. E.DAWSON. 1985. Standards in herpetology andichthyology. Part I. Standard symbolic codes forinstitutional resource collections in herpetology

and ichthyology. Copeia 1985:802–832.SINGH, S. J. 2003. In the Sea of Influence: A World

System Perspective of the Nicobar Islands. LundStudies in Human Ecology 6. Lund University,Lund, Sweden.

TABLE 1. Morphometric comparison of  Calotesandamanensis (ZMUC 36944), and Calotes aurantolabium(BNHM 1436). All distance measurements inmillimeters and mass in grams. Hind limb : SVL 5

ratio of hind limb length to SVL (hind limb 5 sum of femur and tibial lengths). Abbreviations are explainedin text.

Calotes andamanensis Calotes aurantolabium

SVL 84.12 67.88Weight 12.50 6.25HL 23.69 18.20HW 13.62 11.23HW: HL 0.57 0.62Hht 12.42 9.79NW 4.94 4.66TD 2.7 1.00TO 4.99 4.91O 7.39 7.25

Torso 44.42 32.49ShHt 17.28 10.32HpHT 11.17 7.86Humerus 15.44 11.40Ulna 15.33 12.77Femur 19.28 16.10Tibia 19.96 13.79Hind limb : SVL 0.47 0.44FL – 18.09

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SMITH, M. A. 1935. The Fauna of British India,Including Ceylon and Burma: Reptilia and Am-phibia. Vol. II. Sauria. Taylor and Francis, London.

Accepted: 26 January 2008.

APPENDIX 1Specimens Examined

Calotes andamanensis.—ZMUC 36944 (Nicobar Is-lands, India). BNHM 710 (no data).

Calotes aurantolabium.—BNHM 1436 (Kalakad Mun-danthurai Tiger Reserve, Tamil Nadu, India).

Calotes calotes.—UTA 6977, UTA 9382, UTA 9384,UTA 9387, UTA 9383, UTA 9385 (no data).

Calotes ceylonensis.—FMNH 167026, FMNH 167027(Central Province, Sri Lanka).

Calotes liolepis.—UTA 9324 (no data, Sri Lanka).Calotes nemoricola.—FMNH 217712 (Kerala, India).Calotes nigrilabris.—FMNH 131377, FMNH 131378

(Central Province, Sri Lanka); UTA 6979, UTA 9379,UTA 9380, UTA 9381 (no data).

Calotes rouxi.—FMNH 217746 (Kerala, India).Calotes versicolor.—FMNH 191877 (United Provinc-

es, India); ZSI 17399, ZSI 174999, ZSI 21810, ZSI 21423,ZSI 21431, ZSI 21432, ZSI 21434, ZSI 25391, ZSI 25392,ZSI 25398 (no data, India).

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