Amphibia-Reptilia 27 (2006) 1-9

Two new species of Philautus (Anura Ranidae Rhacophorinae)from the Western Ghats India

SD Biju12 Franky Bossuyt1

Abstract The oriental shrubfrog genus Philautus (Anura Ranidae Rhacophorinae) is a large group of direct-developingfrogs that still holds several undescribed species on the Indian subcontinent Here we describe two new species Philautusanili sp nov and Philautus dubois sp nov from Wayanad (Kerala) and Kodaikanal (Tamil Nadu) respectively Both specieswere collected from the township where they are abundant in roadside vegetation and gardens illustrating that several ofthe most common Western Ghats frogs are still not scientifically described The two species are clearly distinct from allknown species but are morphologically similar to each other However Philautus anili differs from P dubois by its largersnout-to-vent length the sharp canthus rostralis and the tibia that is longer than the femur

Introduction

The anuran fauna of the Western Ghats is ex-tremely diverse both in terms of species rich-ness (Dutta 1997 Das 2000 Bossuyt 2002Kuramoto and Joshy 2003) and high-level en-demism (Inger 1999 Biju and Bossuyt 2003Roelants et al 2004) One of the groups thatseems to be very well represented in Asia isthe subfamily Rhacophorinae (Ranidae) whichcontains five genera and about 50 species in In-dia (Frost 2004) Bossuyt and Dubois (2001)recognized 84 valid species names in the genusPhilautus a group of direct developing bush-frogs in that subfamily

Among the Western Ghats species of Phi-lautus several are known only from their orig-inal description Most of them were recordedfrom forest areas and the exploration of thebatrachofauna is still concentrated in this kindof habitat especially in National Parks andWildlife Sanctuaries (eg Ravichandran 1997Krishnamurthy et al 2001 Kuramoto andJoshy 2003) However during our ongoingstudy in the Western Ghats we noticed that

1 - Biology Department Unit of Ecology amp SystematicsVrije Universiteit Brussel (VUB) Pleinlaan 2 B-1050Brussels Belgiume-mail fbossuytvubacbe

2 - Division of Conservation biology Tropical Botanic Gar-den and Research Institute Palode Thiruvananthapu-ram 695 562 Kerala India

many populations of Philautus can be found inurban areas such as hill stations In the presentpaper we describe two new species Philautusanili sp nov and P dubois sp nov that are verycommon in the townships of Wayanad and Ko-daikanal respectively These novelties demon-strate that some species of bushfrogs are notfully dependent on rainforests for their survivaland that also isolated vegetation along the road-side and its vicinity can be extremely valuablefor these frogs

Materials and methods

Measurements and terminology follow Bossuyt and Dubois(2001) The following measurements were taken to thenearest 01 mm using a digital slide-caliper (SVL HL andTL) or a binocular microscope with a micrometer ocularSVL (snout-vent length) HW (head width at the angle ofthe jaws) HL (head length from rear of mandible to tipof snout) MN (distance from rear of mandible to nostril)MFE (distance from rear of mandible to anterior orbitalborder of eye) MBE (distance from rear of mandible toposterior orbital border of eye) SL (snout length from tipof snout to anterior orbital border of eye) EL (eye lengthhorizontal distance between bony orbital borders of eye)IUE (inter upper eyelid width the short distance betweenthe upper eyelids) UEW (maximum upper eyelid width)IN (distance between internal border of nostrils) IFE (in-ternal front of eyes shortest distance between anterior or-bital border of eyes) IBE (internal back of eyes shortestdistance between posterior orbital border of eyes) NS (dis-tance from nostril to tip of snout) EN (distance from nos-tril to anterior orbital border of eye) TYD (largest tympa-num diameter) TYE (distance from posterior orbital bor-der of eye to tympanum) FLL (forelimb length from el-bow to base of outer palmar tubercle) HAL (hand length

copy Koninklijke Brill NV Leiden 2006 Also available online - wwwbrillnl

2 SD Biju F Bossuyt

from base of outer palmar tubercle to tip of third finger)TFL (third finger length from base of first subarticular tu-bercle) FDI-IV (disc width on finger I-IV) FWI-IV (width offinger I-IV at base of disk) TDI-V (disk width on toe I-V)TWI-V (width of toe I-V at base of disk) TL (tibia length)TW (maximum shank width) FL (femur length) FOL (footlength from base of inner metatarsal tubercle to tip of fourthtoe) FTL (length of fourth toe from base of first subarticu-lar tubercle to tip of fourth toe) TFOL (distance from heelto tip of fourth toe) IMT (inner metatarsal tubercle length)and ITL (inner toe length) Drawings of the holotype weremade using a stereomicroscope with a drawing tube

The following institutional abbreviations are used BNHSBombay Natural History Society Bombay MaharashtraIndia TBGRI Tropical Botanic Garden and Research In-stitute Thiruvananthapuram Kerala India The new rha-cophorine species are placed in the genus Philautus becauseof their relatively small snout-to-vent size granular bellythe male having a large subgular pouch which is transparentwhile calling and the female having a relatively low numberof large unpigmented eggs The new species were comparedwith all valid species currently recognized in the genus Phi-lautus (Bossuyt and Dubois 2003 Frost 2004) and espe-cially with the name-bearing types of all Indian species (Ap-pendix)

Philutus anili sp nov (figs 1 and 2)

Diagnosis Philautus anili can be distinguishedfrom all known species in the genus by the com-bination of the following characters (a) medium

adult size (male SVL lt 252 mm) (b) pointedsnout (c) flanks and groin deep brown withlight grey blotches (d) anterior surface of thighsand inner side of tibia light chocolate brownblotches alternated with variable sizes of greypatches (e) ventral side grey and dark brownvermiculated throughout This species is mor-phologically similar to Philautus dubois thesecond species described in this paper How-ever P anili differs from P dubois by its largersize (SVL 241 plusmn 08 mm n = 5 vs 202 plusmn06 mm n = 5 in P dubois) the sharp canthusrostralis (vs rounded in P dubois) and the tibiathat is longer than the thigh (ratio TLFL ratiogt 1 vs TLFL ratio lt 1 in P dubois)

Holotype BNHS 4276 an adult male (SVL238 mm) collected by SD Biju on 28 July1997 at an altitude of 1000 m elevation fromKalpatta approximately 1138primeN 7608primeE inWayanad district Kerala India

Paratypes BNHS 4277 an adult male andBNHS 4280 an adult female collected alongwith the holotype BNHS 4278 BNHS 4279and TBGRI 20020057 three adult males col-lected by SD Biju on 8 August 2001 at an alti-

Figure 1 Holotype of Philautus anili sp nov A Dorsal view B ventral view C lateral view of head D ventral view ofhand E ventral view of foot

Two new species of Philautus 3

Figure 2 Dorsolateral view of Philautus anili holotype in life

tude of 1050 m elevation from Sulthanbetheryapproximately 1137primeN 7617primeE in Wayanaddistrict about 30 km from Kalpatta

Description of the Holotype (all measurementsin mm) Medium-sized Philautus (SVL 238)body slender to slightly elongate (figs 1A 2)head length almost equal to width (HW 90HL 92 MN 75 MFE 58 MBE 26) flatabove outline of snout in dorsal view (fig 1A)and in profile (fig 1C) pointed slightly pro-truding snout length (SL 36) longer than hor-izontal diameter of eye (EL 31) canthus ros-tralis sharp loreal region acutely concave in-terorbital area flat about as wide (IUE 25) asupper eyelid (UEW 24) and slightly wider thaninternarial distance (IN 21) distance betweenanterior margins of eyes (IFE 49) 17 timesin distance between posterior margins of eyes(IBE 83) nostrils oval without lateral flap ofskin closer to tip of snout (NS 10) than to frontof eye (EN 21) pupil oval horizontal tympa-

num (TYD 10) rather distinct rounded 3 timesin eye diameter 14 times distance from tym-panum to eye (TYE 07) (fig 1C) vomerineteeth absent tongue large (70 times 37) cordateemarginate lobes obtuse sparsely granular lin-gual papilla absent supratympanic fold distinctfrom back of eye to shoulder (fig 1C) Fore-limb (FLL 55) shorter than hand (HAL 63TFL 37) relative length of fingers I lt II lt

IV lt III tips of fingers with disks (FDI = 08FWI = 05 FDII = 10 FWII = 04 FDIII =12 FWIII = 05 FDIV = 15 FWIV = 05)with distinct cirummarginal grooves all fingerswith lateral dermal fringe on both edges web-bing absent subarticular tubercles prominentrounded single III2 and IV2 weakly developed(fig 1D) prepollex rather distinct oval sin-gle palmar tubercle oval distinct supernumer-ary tubercles present on all fingers (fig 1D)Hind limbs moderately long heels touch withlimbs folded at right angles to the body tibia52 times longer (TL 116) than wide (TW 21)

4 SD Biju F Bossuyt

longer than thigh (FL 108) length of toe IV(FTL 44) 2 times in distance from base of tar-sus to tip of toe IV (FOL 89) relative lengthof toes I lt II lt III lt V lt IV tips oftoes with disks rather wide compared to toewidth (TDI = 07 TWI = 04 TDII = 08TWII = 05 TDIII = 09 TWIII = 05 TDIV =10 TWIV = 05 TDV = 09 TWV = 05)with a distinct circummarginal groove webbingreduced (fig 1E) dermal fringe along toe Vpresent subarticular tubercles rather prominentrounded simple IV3 and V2 weakly developedinner metatarsal tubercle distinct (IMT 08)oval 2 times in length of toe I (ITL 17) super-numerary tubercle present on all toes (fig 1E)Skin of snout and between eyes shagreened witha few tubercles upper eyelids shagreened withsome prominent granular tubercles side of headshagreened back shagreened with some scat-tered weakly developed tubercles upper part offlanks shagreened with some granular projec-tions lower part of flanks granular dorsal partof forelimb thigh tibia and tarsus shagreenedthroat shagreened to granular chest belly andposterior surface of thighs granular (fig 1B)

Colour in life Dorsum light brown with vari-ous amounts of deep brown irregular patches alight brown strip between the eyes forming a tri-angular grey snout loreal and tympanic regionlight brown a broad dark brown inverted lsquoVrsquoon the back running from the level of the fore-limb and dividing into two unequal ends reach-ing to the level of the hind limbs (figs 1A 2) nu-merous minute blackish brown spots throughoutthe dorsum upper eyelids light brown lateralside of abdomen with dark brown blotches alter-nated with light grey patches flanks and groindeep brown with light grey blotches iris yel-lowish red encircled by a grey ring limbs lightbrown dorsally forelimbs and hind limbs withdark brownish cross bands anterior surface ofthighs and inner side of tibia with light choco-late brown blotches (fig 1B) alternated withvariable sizes of grey patches ventrally greywith variable sized dark brown specks forminga vermiculated belly hands and feet greyish

Colour in alcohol Dorsal coloration and mar-kings almost like in life but faded minute blackspots throughout upper eyelid blackish lorealand tympanic region light grey-brown lateralabdominal area light brown alternated with lightgrey flanks and groin brown with grey blotchesforelimbs light brown hind limbs light brownwith strong dark brown cross bands anteriorsurface of thighs and inner side of tibia withlight brown patches alternated with light greyventrally uniform grey with dark brownish ir-regular spots united into patches (fig 1B)

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductin paratype BNHS 4280 zigzag translucentovary large creamy white eggs about 4 mm indiameter

Etymology The species is named after AnilZachariah in appreciation of his tremendoussupport and assistance to SDB right from thebeginning of his study

Variation Measurements (in mm) of the typeseries are given in table 1 The supernumer-ary tubercles are weakly developed in the threeparatypes (BNHS 4277 BNHS 4279 and TB-GRI 20020057) in comparison with the rest ofthe type series In captivity the frogs go througha considerable range of colour changes whichis reflected in the preserved specimens Freshlycaught specimens have a uniform light tan dor-sum and after captivity they turn to brownishblack (fig 2) The inverted lsquoVrsquo marking on thedorsum can be complete or incomplete

Distribution and Natural History Philautusanili is widely distributed in Wayanad districtIt was first located at Kalpatta near a coffeeplantation The three paratypes were collectedfrom Sulthanbethery in Wayanad district about30 km from Kalpatta Three specimens werecollected from about two meters high on way-side vegetation in Kalpatta town the three oth-ers were found about one meter high on a cof-

Two new species of Philautus 5

Table 1 Morphometric measurements (all in mm range mean and standard deviation) for the type specimens of Philautusanili and P dubois

Philautus anili Philautus dubois

Males (5) Female (1) Males (5) Females (2)

Range Mean SD Range Mean SD Range Mean SD

SVL 232-252 241 08 293 192-208 202 06 251-253 252 01HW 90-93 91 12 118 78-86 82 03 93-101 97 06HL 92-94 93 01 125 74-85 80 05 9-96 93 04IFE 43-52 49 03 60 33-42 38 04 44-47 46 02IBE 80-87 83 03 104 67-75 73 03 83-85 84 00IUE 25-34 29 04 36 26-30 28 02 35-36 36 01UEW 20-27 20 03 31 15-21 18 02 19-21 20 04SL 35-43 39 03 51 30-32 31 01 36-39 38 02EL 30-39 33 03 41 24-28 27 02 27-31 29 03TYD 10-10 10 00 17 09-16 11 03 12-15 14 02FLL 55-58 56 01 67 41-49 43 44 52-58 55 04HAL 61-68 65 03 69 51-63 56 05 72-73 73 01TFL 32-40 36 03 32 27-36 27 03 36-40 38 03TL 116-129 122 05 158 89-93 92 01 118-124 121 04FL 102-120 111 07 147 93-101 98 03 109-118 114 06FOL 89-96 90 03 127 71-83 77 05 95-103 99 06FTL 44-56 51 04 62 36-41 38 02 53-53 53 00IMT 05-09 08 01 10 06-09 07 01 08-09 08 01ITL 17-22 20 02 28 15-19 17 02 22-23 23 01

fee plant This species starts calling in the lateevening ie during or immediately after sunset

Philautus dubois sp nov (figs 3 and 4)

Diagnosis Philautus dubois can be distin-guished from all species in the genus bythe combination of the following characters(a) rather small adult size (male SVL lt

208 mm) (b) tibia longer than foot length(c) supernumerary tubercles well-developed onall toes (d) dorsum and especially flanksprominently granular (e) femur coffee brownwith variable-sized light brown and yellowishbrown blotches (f) ventral side of forelimbcoarsely granular Philautus dubois is morpho-logically most similar to P anili (see diagno-sis of the latter for differences) Some colourmorphs of P dubois could be confused withP tinniens However P dubois differs fromP tinniens in having an elongate body (vs ratherrobust body in P tinniens) and the tibia longerthan the foot length (TL 92 plusmn 01 FOL 77 plusmn05 mm n = 5) (vs tibia shorter than foot

length TL 93 plusmn 01 FOL 101 plusmn 11 n = 5 inP tinniens)

Holotype BNHS 4281 an adult male (SVL205 mm) collected by SD Biju on 17 March2001 at an altitude of 1900 m elevationfrom Kodaikanal 1013primeN 7729primeE in Dindi-gal Anna district Tamil Nadu India

Paratypes BNHS 4282-4285 four adult malesand BNHS 4286-4287 two adult females col-lected along with the holotype

Description of the Holotype (all measurementsin mm) Small-sized Philautus (SVL 205)body slightly elongate (fig 3A) head length al-most equal to width (HW 86 HL 85 MN 73MFE 54 MBE 27) flat above outline of snoutin dorsal view nearly pointed (fig 3A) in pro-file rounded (fig 3B) slightly protruding snoutlength (SL 31) slightly longer than horizon-tal diameter of eye (EL 28) canthus rostralisrounded loreal region acutely concave interor-bital area concave 15 times wider (IUE 27)than upper eyelid (UEW 18) and wider thaninternarial distance (IN 20) distance between

6 SD Biju F Bossuyt

Figure 3 Holotype of Philautus dubois sp nov A Dorsal view B ventral view C lateral view of head D ventral view ofhand E ventral view of foot

anterior margins of eyes (IFE 41) 18 timesin distance between posterior margins of eye(IBE 75) nostrils oval without lateral flapof skin closer to tip of snout (NS 14) thanto front of eye (EN 17) pupil oval tympa-num (TYD 16) rather distinct rounded about17 times in eye diameter (fig 3C) larger thandistance from tympanum to eye (TYE 05)vomerine teeth absent tongue large (45 times29) cordate deeply emarginate lobes obtusesmooth lingual papilla absent supratympanicfold distinct from back of eye to shoulder(fig 3C) Forelimbs (FLL 44) shorter than hand(HAL 63 TFL 31) relative length of fingersI lt II lt IV lt III tips of fingers withdisks (FDI = 07 FWI = 05 FDII = 08FWII = 05 FDIII = 14 FWIII = 06 FDIV =15 FWIV = 06) with distinct cirummar-ginal grooves all fingers without lateral dermal

fringe webbing absent subarticular tuberclesprominent rounded single III2 weakly devel-oped and IV2 absent (fig 3D) prepollex ratherdistinct rounded single palmar tubercle ovaldistinct supernumerary tubercles present on allfingers (fig 3D) Hind limbs moderately longheels barely touch with limbs folded at right an-gles to the body tibia 33 times longer (TL 90)than wide (TW 27) shorter than thigh (FL 94)length of toe IV (FTL 38) 21 times in distancefrom base of tarsus to tip of toe IV (FOL 80)relative length of toes I lt II lt III lt V lt

IV tips of toes with disks rather wide com-pared to toe width (TDI = 07 TWI = 05TDII = 07 TWII = 05 TDIII = 08 TWIII =05 TDIV = 10 TWIV = 07 TDV = 10TWV = 06) with a distinct circummarginalgroove webbing small (fig 3E) dermal fringeor ridge along toe V present subarticular tuber-

Two new species of Philautus 7

Figure 4 Life specimens of Philautus dubois from the type locality A Paratype (BNHS 4286 female) dorsolateral viewB Paratype (BNHS 4287 female) dorsolateral view C Male lateral view (not captured) D male dorsal view (not captured)

cles rather prominent rounded simple IV 23 and V2 weakly developed inner metatarsaltubercle distinct (IMT 09) oval more than2 times in length of toe I (ITL 19) supernu-merary tubercle present on all toes (fig 3E)Skin of snout and between eyes shagreened up-per eyelids slightly granular side of head espe-cially tympanic region granular (fig 3C) backand flanks granular dorsal part of forelimbsshagreened to sparsely granular thigh and tibiagranular tarsus shagreened ventral part of fore-limbs granular throat shagreened to granularchest belly and posterior surface of thighs gran-ular (fig 3B)

Colour in life Dorsally pale grayish greentinged with shining silver and cobalt violet adark grey inverted lsquoVrsquo on the dorsum runningfrom the level of the eye to nearly the level

of the vent (fig 3A) with black brown andsilver violet spots of variable size present es-pecially on flanks loreal and temporal regionbrownish black iris golden brown with a green-ish brown outer ring pupil with a golden outerring limbs light grey-brown with blackish crossbands finger tips white femur coffee brownwith variable-sized light brown and yellowishbrown blotches tibia and tarsus dark brownwith grayish cross bands toe discs white ven-trally with variable amounts of dark and lightpatches and spots side of the belly with lightyellowish brown blotches ventral surface ofthigh tibia and tarsus grayish brown intermin-gled with brown blotches feet and hands darkgrayish brown with black spots

Colour in alcohol Dorsum light grayishbrown black and grey spots throughout in-

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

Two new species of Philautus 3

Figure 2 Dorsolateral view of Philautus anili holotype in life

tude of 1050 m elevation from Sulthanbetheryapproximately 1137primeN 7617primeE in Wayanaddistrict about 30 km from Kalpatta

Description of the Holotype (all measurementsin mm) Medium-sized Philautus (SVL 238)body slender to slightly elongate (figs 1A 2)head length almost equal to width (HW 90HL 92 MN 75 MFE 58 MBE 26) flatabove outline of snout in dorsal view (fig 1A)and in profile (fig 1C) pointed slightly pro-truding snout length (SL 36) longer than hor-izontal diameter of eye (EL 31) canthus ros-tralis sharp loreal region acutely concave in-terorbital area flat about as wide (IUE 25) asupper eyelid (UEW 24) and slightly wider thaninternarial distance (IN 21) distance betweenanterior margins of eyes (IFE 49) 17 timesin distance between posterior margins of eyes(IBE 83) nostrils oval without lateral flap ofskin closer to tip of snout (NS 10) than to frontof eye (EN 21) pupil oval horizontal tympa-

num (TYD 10) rather distinct rounded 3 timesin eye diameter 14 times distance from tym-panum to eye (TYE 07) (fig 1C) vomerineteeth absent tongue large (70 times 37) cordateemarginate lobes obtuse sparsely granular lin-gual papilla absent supratympanic fold distinctfrom back of eye to shoulder (fig 1C) Fore-limb (FLL 55) shorter than hand (HAL 63TFL 37) relative length of fingers I lt II lt

IV lt III tips of fingers with disks (FDI = 08FWI = 05 FDII = 10 FWII = 04 FDIII =12 FWIII = 05 FDIV = 15 FWIV = 05)with distinct cirummarginal grooves all fingerswith lateral dermal fringe on both edges web-bing absent subarticular tubercles prominentrounded single III2 and IV2 weakly developed(fig 1D) prepollex rather distinct oval sin-gle palmar tubercle oval distinct supernumer-ary tubercles present on all fingers (fig 1D)Hind limbs moderately long heels touch withlimbs folded at right angles to the body tibia52 times longer (TL 116) than wide (TW 21)

4 SD Biju F Bossuyt

longer than thigh (FL 108) length of toe IV(FTL 44) 2 times in distance from base of tar-sus to tip of toe IV (FOL 89) relative lengthof toes I lt II lt III lt V lt IV tips oftoes with disks rather wide compared to toewidth (TDI = 07 TWI = 04 TDII = 08TWII = 05 TDIII = 09 TWIII = 05 TDIV =10 TWIV = 05 TDV = 09 TWV = 05)with a distinct circummarginal groove webbingreduced (fig 1E) dermal fringe along toe Vpresent subarticular tubercles rather prominentrounded simple IV3 and V2 weakly developedinner metatarsal tubercle distinct (IMT 08)oval 2 times in length of toe I (ITL 17) super-numerary tubercle present on all toes (fig 1E)Skin of snout and between eyes shagreened witha few tubercles upper eyelids shagreened withsome prominent granular tubercles side of headshagreened back shagreened with some scat-tered weakly developed tubercles upper part offlanks shagreened with some granular projec-tions lower part of flanks granular dorsal partof forelimb thigh tibia and tarsus shagreenedthroat shagreened to granular chest belly andposterior surface of thighs granular (fig 1B)

Colour in life Dorsum light brown with vari-ous amounts of deep brown irregular patches alight brown strip between the eyes forming a tri-angular grey snout loreal and tympanic regionlight brown a broad dark brown inverted lsquoVrsquoon the back running from the level of the fore-limb and dividing into two unequal ends reach-ing to the level of the hind limbs (figs 1A 2) nu-merous minute blackish brown spots throughoutthe dorsum upper eyelids light brown lateralside of abdomen with dark brown blotches alter-nated with light grey patches flanks and groindeep brown with light grey blotches iris yel-lowish red encircled by a grey ring limbs lightbrown dorsally forelimbs and hind limbs withdark brownish cross bands anterior surface ofthighs and inner side of tibia with light choco-late brown blotches (fig 1B) alternated withvariable sizes of grey patches ventrally greywith variable sized dark brown specks forminga vermiculated belly hands and feet greyish

Colour in alcohol Dorsal coloration and mar-kings almost like in life but faded minute blackspots throughout upper eyelid blackish lorealand tympanic region light grey-brown lateralabdominal area light brown alternated with lightgrey flanks and groin brown with grey blotchesforelimbs light brown hind limbs light brownwith strong dark brown cross bands anteriorsurface of thighs and inner side of tibia withlight brown patches alternated with light greyventrally uniform grey with dark brownish ir-regular spots united into patches (fig 1B)

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductin paratype BNHS 4280 zigzag translucentovary large creamy white eggs about 4 mm indiameter

Etymology The species is named after AnilZachariah in appreciation of his tremendoussupport and assistance to SDB right from thebeginning of his study

Variation Measurements (in mm) of the typeseries are given in table 1 The supernumer-ary tubercles are weakly developed in the threeparatypes (BNHS 4277 BNHS 4279 and TB-GRI 20020057) in comparison with the rest ofthe type series In captivity the frogs go througha considerable range of colour changes whichis reflected in the preserved specimens Freshlycaught specimens have a uniform light tan dor-sum and after captivity they turn to brownishblack (fig 2) The inverted lsquoVrsquo marking on thedorsum can be complete or incomplete

Distribution and Natural History Philautusanili is widely distributed in Wayanad districtIt was first located at Kalpatta near a coffeeplantation The three paratypes were collectedfrom Sulthanbethery in Wayanad district about30 km from Kalpatta Three specimens werecollected from about two meters high on way-side vegetation in Kalpatta town the three oth-ers were found about one meter high on a cof-

Two new species of Philautus 5

Table 1 Morphometric measurements (all in mm range mean and standard deviation) for the type specimens of Philautusanili and P dubois

Philautus anili Philautus dubois

Males (5) Female (1) Males (5) Females (2)

Range Mean SD Range Mean SD Range Mean SD

SVL 232-252 241 08 293 192-208 202 06 251-253 252 01HW 90-93 91 12 118 78-86 82 03 93-101 97 06HL 92-94 93 01 125 74-85 80 05 9-96 93 04IFE 43-52 49 03 60 33-42 38 04 44-47 46 02IBE 80-87 83 03 104 67-75 73 03 83-85 84 00IUE 25-34 29 04 36 26-30 28 02 35-36 36 01UEW 20-27 20 03 31 15-21 18 02 19-21 20 04SL 35-43 39 03 51 30-32 31 01 36-39 38 02EL 30-39 33 03 41 24-28 27 02 27-31 29 03TYD 10-10 10 00 17 09-16 11 03 12-15 14 02FLL 55-58 56 01 67 41-49 43 44 52-58 55 04HAL 61-68 65 03 69 51-63 56 05 72-73 73 01TFL 32-40 36 03 32 27-36 27 03 36-40 38 03TL 116-129 122 05 158 89-93 92 01 118-124 121 04FL 102-120 111 07 147 93-101 98 03 109-118 114 06FOL 89-96 90 03 127 71-83 77 05 95-103 99 06FTL 44-56 51 04 62 36-41 38 02 53-53 53 00IMT 05-09 08 01 10 06-09 07 01 08-09 08 01ITL 17-22 20 02 28 15-19 17 02 22-23 23 01

fee plant This species starts calling in the lateevening ie during or immediately after sunset

Philautus dubois sp nov (figs 3 and 4)

Diagnosis Philautus dubois can be distin-guished from all species in the genus bythe combination of the following characters(a) rather small adult size (male SVL lt

208 mm) (b) tibia longer than foot length(c) supernumerary tubercles well-developed onall toes (d) dorsum and especially flanksprominently granular (e) femur coffee brownwith variable-sized light brown and yellowishbrown blotches (f) ventral side of forelimbcoarsely granular Philautus dubois is morpho-logically most similar to P anili (see diagno-sis of the latter for differences) Some colourmorphs of P dubois could be confused withP tinniens However P dubois differs fromP tinniens in having an elongate body (vs ratherrobust body in P tinniens) and the tibia longerthan the foot length (TL 92 plusmn 01 FOL 77 plusmn05 mm n = 5) (vs tibia shorter than foot

length TL 93 plusmn 01 FOL 101 plusmn 11 n = 5 inP tinniens)

Holotype BNHS 4281 an adult male (SVL205 mm) collected by SD Biju on 17 March2001 at an altitude of 1900 m elevationfrom Kodaikanal 1013primeN 7729primeE in Dindi-gal Anna district Tamil Nadu India

Paratypes BNHS 4282-4285 four adult malesand BNHS 4286-4287 two adult females col-lected along with the holotype

Description of the Holotype (all measurementsin mm) Small-sized Philautus (SVL 205)body slightly elongate (fig 3A) head length al-most equal to width (HW 86 HL 85 MN 73MFE 54 MBE 27) flat above outline of snoutin dorsal view nearly pointed (fig 3A) in pro-file rounded (fig 3B) slightly protruding snoutlength (SL 31) slightly longer than horizon-tal diameter of eye (EL 28) canthus rostralisrounded loreal region acutely concave interor-bital area concave 15 times wider (IUE 27)than upper eyelid (UEW 18) and wider thaninternarial distance (IN 20) distance between

6 SD Biju F Bossuyt

Figure 3 Holotype of Philautus dubois sp nov A Dorsal view B ventral view C lateral view of head D ventral view ofhand E ventral view of foot

anterior margins of eyes (IFE 41) 18 timesin distance between posterior margins of eye(IBE 75) nostrils oval without lateral flapof skin closer to tip of snout (NS 14) thanto front of eye (EN 17) pupil oval tympa-num (TYD 16) rather distinct rounded about17 times in eye diameter (fig 3C) larger thandistance from tympanum to eye (TYE 05)vomerine teeth absent tongue large (45 times29) cordate deeply emarginate lobes obtusesmooth lingual papilla absent supratympanicfold distinct from back of eye to shoulder(fig 3C) Forelimbs (FLL 44) shorter than hand(HAL 63 TFL 31) relative length of fingersI lt II lt IV lt III tips of fingers withdisks (FDI = 07 FWI = 05 FDII = 08FWII = 05 FDIII = 14 FWIII = 06 FDIV =15 FWIV = 06) with distinct cirummar-ginal grooves all fingers without lateral dermal

fringe webbing absent subarticular tuberclesprominent rounded single III2 weakly devel-oped and IV2 absent (fig 3D) prepollex ratherdistinct rounded single palmar tubercle ovaldistinct supernumerary tubercles present on allfingers (fig 3D) Hind limbs moderately longheels barely touch with limbs folded at right an-gles to the body tibia 33 times longer (TL 90)than wide (TW 27) shorter than thigh (FL 94)length of toe IV (FTL 38) 21 times in distancefrom base of tarsus to tip of toe IV (FOL 80)relative length of toes I lt II lt III lt V lt

IV tips of toes with disks rather wide com-pared to toe width (TDI = 07 TWI = 05TDII = 07 TWII = 05 TDIII = 08 TWIII =05 TDIV = 10 TWIV = 07 TDV = 10TWV = 06) with a distinct circummarginalgroove webbing small (fig 3E) dermal fringeor ridge along toe V present subarticular tuber-

Two new species of Philautus 7

Figure 4 Life specimens of Philautus dubois from the type locality A Paratype (BNHS 4286 female) dorsolateral viewB Paratype (BNHS 4287 female) dorsolateral view C Male lateral view (not captured) D male dorsal view (not captured)

cles rather prominent rounded simple IV 23 and V2 weakly developed inner metatarsaltubercle distinct (IMT 09) oval more than2 times in length of toe I (ITL 19) supernu-merary tubercle present on all toes (fig 3E)Skin of snout and between eyes shagreened up-per eyelids slightly granular side of head espe-cially tympanic region granular (fig 3C) backand flanks granular dorsal part of forelimbsshagreened to sparsely granular thigh and tibiagranular tarsus shagreened ventral part of fore-limbs granular throat shagreened to granularchest belly and posterior surface of thighs gran-ular (fig 3B)

Colour in life Dorsally pale grayish greentinged with shining silver and cobalt violet adark grey inverted lsquoVrsquo on the dorsum runningfrom the level of the eye to nearly the level

of the vent (fig 3A) with black brown andsilver violet spots of variable size present es-pecially on flanks loreal and temporal regionbrownish black iris golden brown with a green-ish brown outer ring pupil with a golden outerring limbs light grey-brown with blackish crossbands finger tips white femur coffee brownwith variable-sized light brown and yellowishbrown blotches tibia and tarsus dark brownwith grayish cross bands toe discs white ven-trally with variable amounts of dark and lightpatches and spots side of the belly with lightyellowish brown blotches ventral surface ofthigh tibia and tarsus grayish brown intermin-gled with brown blotches feet and hands darkgrayish brown with black spots

Colour in alcohol Dorsum light grayishbrown black and grey spots throughout in-

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

4 SD Biju F Bossuyt

longer than thigh (FL 108) length of toe IV(FTL 44) 2 times in distance from base of tar-sus to tip of toe IV (FOL 89) relative lengthof toes I lt II lt III lt V lt IV tips oftoes with disks rather wide compared to toewidth (TDI = 07 TWI = 04 TDII = 08TWII = 05 TDIII = 09 TWIII = 05 TDIV =10 TWIV = 05 TDV = 09 TWV = 05)with a distinct circummarginal groove webbingreduced (fig 1E) dermal fringe along toe Vpresent subarticular tubercles rather prominentrounded simple IV3 and V2 weakly developedinner metatarsal tubercle distinct (IMT 08)oval 2 times in length of toe I (ITL 17) super-numerary tubercle present on all toes (fig 1E)Skin of snout and between eyes shagreened witha few tubercles upper eyelids shagreened withsome prominent granular tubercles side of headshagreened back shagreened with some scat-tered weakly developed tubercles upper part offlanks shagreened with some granular projec-tions lower part of flanks granular dorsal partof forelimb thigh tibia and tarsus shagreenedthroat shagreened to granular chest belly andposterior surface of thighs granular (fig 1B)

Colour in life Dorsum light brown with vari-ous amounts of deep brown irregular patches alight brown strip between the eyes forming a tri-angular grey snout loreal and tympanic regionlight brown a broad dark brown inverted lsquoVrsquoon the back running from the level of the fore-limb and dividing into two unequal ends reach-ing to the level of the hind limbs (figs 1A 2) nu-merous minute blackish brown spots throughoutthe dorsum upper eyelids light brown lateralside of abdomen with dark brown blotches alter-nated with light grey patches flanks and groindeep brown with light grey blotches iris yel-lowish red encircled by a grey ring limbs lightbrown dorsally forelimbs and hind limbs withdark brownish cross bands anterior surface ofthighs and inner side of tibia with light choco-late brown blotches (fig 1B) alternated withvariable sizes of grey patches ventrally greywith variable sized dark brown specks forminga vermiculated belly hands and feet greyish

Colour in alcohol Dorsal coloration and mar-kings almost like in life but faded minute blackspots throughout upper eyelid blackish lorealand tympanic region light grey-brown lateralabdominal area light brown alternated with lightgrey flanks and groin brown with grey blotchesforelimbs light brown hind limbs light brownwith strong dark brown cross bands anteriorsurface of thighs and inner side of tibia withlight brown patches alternated with light greyventrally uniform grey with dark brownish ir-regular spots united into patches (fig 1B)

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductin paratype BNHS 4280 zigzag translucentovary large creamy white eggs about 4 mm indiameter

Etymology The species is named after AnilZachariah in appreciation of his tremendoussupport and assistance to SDB right from thebeginning of his study

Variation Measurements (in mm) of the typeseries are given in table 1 The supernumer-ary tubercles are weakly developed in the threeparatypes (BNHS 4277 BNHS 4279 and TB-GRI 20020057) in comparison with the rest ofthe type series In captivity the frogs go througha considerable range of colour changes whichis reflected in the preserved specimens Freshlycaught specimens have a uniform light tan dor-sum and after captivity they turn to brownishblack (fig 2) The inverted lsquoVrsquo marking on thedorsum can be complete or incomplete

Distribution and Natural History Philautusanili is widely distributed in Wayanad districtIt was first located at Kalpatta near a coffeeplantation The three paratypes were collectedfrom Sulthanbethery in Wayanad district about30 km from Kalpatta Three specimens werecollected from about two meters high on way-side vegetation in Kalpatta town the three oth-ers were found about one meter high on a cof-

Two new species of Philautus 5

Table 1 Morphometric measurements (all in mm range mean and standard deviation) for the type specimens of Philautusanili and P dubois

Philautus anili Philautus dubois

Males (5) Female (1) Males (5) Females (2)

Range Mean SD Range Mean SD Range Mean SD

SVL 232-252 241 08 293 192-208 202 06 251-253 252 01HW 90-93 91 12 118 78-86 82 03 93-101 97 06HL 92-94 93 01 125 74-85 80 05 9-96 93 04IFE 43-52 49 03 60 33-42 38 04 44-47 46 02IBE 80-87 83 03 104 67-75 73 03 83-85 84 00IUE 25-34 29 04 36 26-30 28 02 35-36 36 01UEW 20-27 20 03 31 15-21 18 02 19-21 20 04SL 35-43 39 03 51 30-32 31 01 36-39 38 02EL 30-39 33 03 41 24-28 27 02 27-31 29 03TYD 10-10 10 00 17 09-16 11 03 12-15 14 02FLL 55-58 56 01 67 41-49 43 44 52-58 55 04HAL 61-68 65 03 69 51-63 56 05 72-73 73 01TFL 32-40 36 03 32 27-36 27 03 36-40 38 03TL 116-129 122 05 158 89-93 92 01 118-124 121 04FL 102-120 111 07 147 93-101 98 03 109-118 114 06FOL 89-96 90 03 127 71-83 77 05 95-103 99 06FTL 44-56 51 04 62 36-41 38 02 53-53 53 00IMT 05-09 08 01 10 06-09 07 01 08-09 08 01ITL 17-22 20 02 28 15-19 17 02 22-23 23 01

fee plant This species starts calling in the lateevening ie during or immediately after sunset

Philautus dubois sp nov (figs 3 and 4)

Diagnosis Philautus dubois can be distin-guished from all species in the genus bythe combination of the following characters(a) rather small adult size (male SVL lt

208 mm) (b) tibia longer than foot length(c) supernumerary tubercles well-developed onall toes (d) dorsum and especially flanksprominently granular (e) femur coffee brownwith variable-sized light brown and yellowishbrown blotches (f) ventral side of forelimbcoarsely granular Philautus dubois is morpho-logically most similar to P anili (see diagno-sis of the latter for differences) Some colourmorphs of P dubois could be confused withP tinniens However P dubois differs fromP tinniens in having an elongate body (vs ratherrobust body in P tinniens) and the tibia longerthan the foot length (TL 92 plusmn 01 FOL 77 plusmn05 mm n = 5) (vs tibia shorter than foot

length TL 93 plusmn 01 FOL 101 plusmn 11 n = 5 inP tinniens)

Holotype BNHS 4281 an adult male (SVL205 mm) collected by SD Biju on 17 March2001 at an altitude of 1900 m elevationfrom Kodaikanal 1013primeN 7729primeE in Dindi-gal Anna district Tamil Nadu India

Paratypes BNHS 4282-4285 four adult malesand BNHS 4286-4287 two adult females col-lected along with the holotype

Description of the Holotype (all measurementsin mm) Small-sized Philautus (SVL 205)body slightly elongate (fig 3A) head length al-most equal to width (HW 86 HL 85 MN 73MFE 54 MBE 27) flat above outline of snoutin dorsal view nearly pointed (fig 3A) in pro-file rounded (fig 3B) slightly protruding snoutlength (SL 31) slightly longer than horizon-tal diameter of eye (EL 28) canthus rostralisrounded loreal region acutely concave interor-bital area concave 15 times wider (IUE 27)than upper eyelid (UEW 18) and wider thaninternarial distance (IN 20) distance between

6 SD Biju F Bossuyt

Figure 3 Holotype of Philautus dubois sp nov A Dorsal view B ventral view C lateral view of head D ventral view ofhand E ventral view of foot

anterior margins of eyes (IFE 41) 18 timesin distance between posterior margins of eye(IBE 75) nostrils oval without lateral flapof skin closer to tip of snout (NS 14) thanto front of eye (EN 17) pupil oval tympa-num (TYD 16) rather distinct rounded about17 times in eye diameter (fig 3C) larger thandistance from tympanum to eye (TYE 05)vomerine teeth absent tongue large (45 times29) cordate deeply emarginate lobes obtusesmooth lingual papilla absent supratympanicfold distinct from back of eye to shoulder(fig 3C) Forelimbs (FLL 44) shorter than hand(HAL 63 TFL 31) relative length of fingersI lt II lt IV lt III tips of fingers withdisks (FDI = 07 FWI = 05 FDII = 08FWII = 05 FDIII = 14 FWIII = 06 FDIV =15 FWIV = 06) with distinct cirummar-ginal grooves all fingers without lateral dermal

fringe webbing absent subarticular tuberclesprominent rounded single III2 weakly devel-oped and IV2 absent (fig 3D) prepollex ratherdistinct rounded single palmar tubercle ovaldistinct supernumerary tubercles present on allfingers (fig 3D) Hind limbs moderately longheels barely touch with limbs folded at right an-gles to the body tibia 33 times longer (TL 90)than wide (TW 27) shorter than thigh (FL 94)length of toe IV (FTL 38) 21 times in distancefrom base of tarsus to tip of toe IV (FOL 80)relative length of toes I lt II lt III lt V lt

IV tips of toes with disks rather wide com-pared to toe width (TDI = 07 TWI = 05TDII = 07 TWII = 05 TDIII = 08 TWIII =05 TDIV = 10 TWIV = 07 TDV = 10TWV = 06) with a distinct circummarginalgroove webbing small (fig 3E) dermal fringeor ridge along toe V present subarticular tuber-

Two new species of Philautus 7

Figure 4 Life specimens of Philautus dubois from the type locality A Paratype (BNHS 4286 female) dorsolateral viewB Paratype (BNHS 4287 female) dorsolateral view C Male lateral view (not captured) D male dorsal view (not captured)

cles rather prominent rounded simple IV 23 and V2 weakly developed inner metatarsaltubercle distinct (IMT 09) oval more than2 times in length of toe I (ITL 19) supernu-merary tubercle present on all toes (fig 3E)Skin of snout and between eyes shagreened up-per eyelids slightly granular side of head espe-cially tympanic region granular (fig 3C) backand flanks granular dorsal part of forelimbsshagreened to sparsely granular thigh and tibiagranular tarsus shagreened ventral part of fore-limbs granular throat shagreened to granularchest belly and posterior surface of thighs gran-ular (fig 3B)

Colour in life Dorsally pale grayish greentinged with shining silver and cobalt violet adark grey inverted lsquoVrsquo on the dorsum runningfrom the level of the eye to nearly the level

of the vent (fig 3A) with black brown andsilver violet spots of variable size present es-pecially on flanks loreal and temporal regionbrownish black iris golden brown with a green-ish brown outer ring pupil with a golden outerring limbs light grey-brown with blackish crossbands finger tips white femur coffee brownwith variable-sized light brown and yellowishbrown blotches tibia and tarsus dark brownwith grayish cross bands toe discs white ven-trally with variable amounts of dark and lightpatches and spots side of the belly with lightyellowish brown blotches ventral surface ofthigh tibia and tarsus grayish brown intermin-gled with brown blotches feet and hands darkgrayish brown with black spots

Colour in alcohol Dorsum light grayishbrown black and grey spots throughout in-

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

Two new species of Philautus 5

Table 1 Morphometric measurements (all in mm range mean and standard deviation) for the type specimens of Philautusanili and P dubois

Philautus anili Philautus dubois

Males (5) Female (1) Males (5) Females (2)

Range Mean SD Range Mean SD Range Mean SD

SVL 232-252 241 08 293 192-208 202 06 251-253 252 01HW 90-93 91 12 118 78-86 82 03 93-101 97 06HL 92-94 93 01 125 74-85 80 05 9-96 93 04IFE 43-52 49 03 60 33-42 38 04 44-47 46 02IBE 80-87 83 03 104 67-75 73 03 83-85 84 00IUE 25-34 29 04 36 26-30 28 02 35-36 36 01UEW 20-27 20 03 31 15-21 18 02 19-21 20 04SL 35-43 39 03 51 30-32 31 01 36-39 38 02EL 30-39 33 03 41 24-28 27 02 27-31 29 03TYD 10-10 10 00 17 09-16 11 03 12-15 14 02FLL 55-58 56 01 67 41-49 43 44 52-58 55 04HAL 61-68 65 03 69 51-63 56 05 72-73 73 01TFL 32-40 36 03 32 27-36 27 03 36-40 38 03TL 116-129 122 05 158 89-93 92 01 118-124 121 04FL 102-120 111 07 147 93-101 98 03 109-118 114 06FOL 89-96 90 03 127 71-83 77 05 95-103 99 06FTL 44-56 51 04 62 36-41 38 02 53-53 53 00IMT 05-09 08 01 10 06-09 07 01 08-09 08 01ITL 17-22 20 02 28 15-19 17 02 22-23 23 01

fee plant This species starts calling in the lateevening ie during or immediately after sunset

Philautus dubois sp nov (figs 3 and 4)

Diagnosis Philautus dubois can be distin-guished from all species in the genus bythe combination of the following characters(a) rather small adult size (male SVL lt

208 mm) (b) tibia longer than foot length(c) supernumerary tubercles well-developed onall toes (d) dorsum and especially flanksprominently granular (e) femur coffee brownwith variable-sized light brown and yellowishbrown blotches (f) ventral side of forelimbcoarsely granular Philautus dubois is morpho-logically most similar to P anili (see diagno-sis of the latter for differences) Some colourmorphs of P dubois could be confused withP tinniens However P dubois differs fromP tinniens in having an elongate body (vs ratherrobust body in P tinniens) and the tibia longerthan the foot length (TL 92 plusmn 01 FOL 77 plusmn05 mm n = 5) (vs tibia shorter than foot

length TL 93 plusmn 01 FOL 101 plusmn 11 n = 5 inP tinniens)

Holotype BNHS 4281 an adult male (SVL205 mm) collected by SD Biju on 17 March2001 at an altitude of 1900 m elevationfrom Kodaikanal 1013primeN 7729primeE in Dindi-gal Anna district Tamil Nadu India

Paratypes BNHS 4282-4285 four adult malesand BNHS 4286-4287 two adult females col-lected along with the holotype

Description of the Holotype (all measurementsin mm) Small-sized Philautus (SVL 205)body slightly elongate (fig 3A) head length al-most equal to width (HW 86 HL 85 MN 73MFE 54 MBE 27) flat above outline of snoutin dorsal view nearly pointed (fig 3A) in pro-file rounded (fig 3B) slightly protruding snoutlength (SL 31) slightly longer than horizon-tal diameter of eye (EL 28) canthus rostralisrounded loreal region acutely concave interor-bital area concave 15 times wider (IUE 27)than upper eyelid (UEW 18) and wider thaninternarial distance (IN 20) distance between

6 SD Biju F Bossuyt

Figure 3 Holotype of Philautus dubois sp nov A Dorsal view B ventral view C lateral view of head D ventral view ofhand E ventral view of foot

anterior margins of eyes (IFE 41) 18 timesin distance between posterior margins of eye(IBE 75) nostrils oval without lateral flapof skin closer to tip of snout (NS 14) thanto front of eye (EN 17) pupil oval tympa-num (TYD 16) rather distinct rounded about17 times in eye diameter (fig 3C) larger thandistance from tympanum to eye (TYE 05)vomerine teeth absent tongue large (45 times29) cordate deeply emarginate lobes obtusesmooth lingual papilla absent supratympanicfold distinct from back of eye to shoulder(fig 3C) Forelimbs (FLL 44) shorter than hand(HAL 63 TFL 31) relative length of fingersI lt II lt IV lt III tips of fingers withdisks (FDI = 07 FWI = 05 FDII = 08FWII = 05 FDIII = 14 FWIII = 06 FDIV =15 FWIV = 06) with distinct cirummar-ginal grooves all fingers without lateral dermal

fringe webbing absent subarticular tuberclesprominent rounded single III2 weakly devel-oped and IV2 absent (fig 3D) prepollex ratherdistinct rounded single palmar tubercle ovaldistinct supernumerary tubercles present on allfingers (fig 3D) Hind limbs moderately longheels barely touch with limbs folded at right an-gles to the body tibia 33 times longer (TL 90)than wide (TW 27) shorter than thigh (FL 94)length of toe IV (FTL 38) 21 times in distancefrom base of tarsus to tip of toe IV (FOL 80)relative length of toes I lt II lt III lt V lt

IV tips of toes with disks rather wide com-pared to toe width (TDI = 07 TWI = 05TDII = 07 TWII = 05 TDIII = 08 TWIII =05 TDIV = 10 TWIV = 07 TDV = 10TWV = 06) with a distinct circummarginalgroove webbing small (fig 3E) dermal fringeor ridge along toe V present subarticular tuber-

Two new species of Philautus 7

Figure 4 Life specimens of Philautus dubois from the type locality A Paratype (BNHS 4286 female) dorsolateral viewB Paratype (BNHS 4287 female) dorsolateral view C Male lateral view (not captured) D male dorsal view (not captured)

cles rather prominent rounded simple IV 23 and V2 weakly developed inner metatarsaltubercle distinct (IMT 09) oval more than2 times in length of toe I (ITL 19) supernu-merary tubercle present on all toes (fig 3E)Skin of snout and between eyes shagreened up-per eyelids slightly granular side of head espe-cially tympanic region granular (fig 3C) backand flanks granular dorsal part of forelimbsshagreened to sparsely granular thigh and tibiagranular tarsus shagreened ventral part of fore-limbs granular throat shagreened to granularchest belly and posterior surface of thighs gran-ular (fig 3B)

Colour in life Dorsally pale grayish greentinged with shining silver and cobalt violet adark grey inverted lsquoVrsquo on the dorsum runningfrom the level of the eye to nearly the level

of the vent (fig 3A) with black brown andsilver violet spots of variable size present es-pecially on flanks loreal and temporal regionbrownish black iris golden brown with a green-ish brown outer ring pupil with a golden outerring limbs light grey-brown with blackish crossbands finger tips white femur coffee brownwith variable-sized light brown and yellowishbrown blotches tibia and tarsus dark brownwith grayish cross bands toe discs white ven-trally with variable amounts of dark and lightpatches and spots side of the belly with lightyellowish brown blotches ventral surface ofthigh tibia and tarsus grayish brown intermin-gled with brown blotches feet and hands darkgrayish brown with black spots

Colour in alcohol Dorsum light grayishbrown black and grey spots throughout in-

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

6 SD Biju F Bossuyt

Figure 3 Holotype of Philautus dubois sp nov A Dorsal view B ventral view C lateral view of head D ventral view ofhand E ventral view of foot

anterior margins of eyes (IFE 41) 18 timesin distance between posterior margins of eye(IBE 75) nostrils oval without lateral flapof skin closer to tip of snout (NS 14) thanto front of eye (EN 17) pupil oval tympa-num (TYD 16) rather distinct rounded about17 times in eye diameter (fig 3C) larger thandistance from tympanum to eye (TYE 05)vomerine teeth absent tongue large (45 times29) cordate deeply emarginate lobes obtusesmooth lingual papilla absent supratympanicfold distinct from back of eye to shoulder(fig 3C) Forelimbs (FLL 44) shorter than hand(HAL 63 TFL 31) relative length of fingersI lt II lt IV lt III tips of fingers withdisks (FDI = 07 FWI = 05 FDII = 08FWII = 05 FDIII = 14 FWIII = 06 FDIV =15 FWIV = 06) with distinct cirummar-ginal grooves all fingers without lateral dermal

fringe webbing absent subarticular tuberclesprominent rounded single III2 weakly devel-oped and IV2 absent (fig 3D) prepollex ratherdistinct rounded single palmar tubercle ovaldistinct supernumerary tubercles present on allfingers (fig 3D) Hind limbs moderately longheels barely touch with limbs folded at right an-gles to the body tibia 33 times longer (TL 90)than wide (TW 27) shorter than thigh (FL 94)length of toe IV (FTL 38) 21 times in distancefrom base of tarsus to tip of toe IV (FOL 80)relative length of toes I lt II lt III lt V lt

IV tips of toes with disks rather wide com-pared to toe width (TDI = 07 TWI = 05TDII = 07 TWII = 05 TDIII = 08 TWIII =05 TDIV = 10 TWIV = 07 TDV = 10TWV = 06) with a distinct circummarginalgroove webbing small (fig 3E) dermal fringeor ridge along toe V present subarticular tuber-

Two new species of Philautus 7

Figure 4 Life specimens of Philautus dubois from the type locality A Paratype (BNHS 4286 female) dorsolateral viewB Paratype (BNHS 4287 female) dorsolateral view C Male lateral view (not captured) D male dorsal view (not captured)

cles rather prominent rounded simple IV 23 and V2 weakly developed inner metatarsaltubercle distinct (IMT 09) oval more than2 times in length of toe I (ITL 19) supernu-merary tubercle present on all toes (fig 3E)Skin of snout and between eyes shagreened up-per eyelids slightly granular side of head espe-cially tympanic region granular (fig 3C) backand flanks granular dorsal part of forelimbsshagreened to sparsely granular thigh and tibiagranular tarsus shagreened ventral part of fore-limbs granular throat shagreened to granularchest belly and posterior surface of thighs gran-ular (fig 3B)

Colour in life Dorsally pale grayish greentinged with shining silver and cobalt violet adark grey inverted lsquoVrsquo on the dorsum runningfrom the level of the eye to nearly the level

of the vent (fig 3A) with black brown andsilver violet spots of variable size present es-pecially on flanks loreal and temporal regionbrownish black iris golden brown with a green-ish brown outer ring pupil with a golden outerring limbs light grey-brown with blackish crossbands finger tips white femur coffee brownwith variable-sized light brown and yellowishbrown blotches tibia and tarsus dark brownwith grayish cross bands toe discs white ven-trally with variable amounts of dark and lightpatches and spots side of the belly with lightyellowish brown blotches ventral surface ofthigh tibia and tarsus grayish brown intermin-gled with brown blotches feet and hands darkgrayish brown with black spots

Colour in alcohol Dorsum light grayishbrown black and grey spots throughout in-

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

Two new species of Philautus 7

Figure 4 Life specimens of Philautus dubois from the type locality A Paratype (BNHS 4286 female) dorsolateral viewB Paratype (BNHS 4287 female) dorsolateral view C Male lateral view (not captured) D male dorsal view (not captured)

cles rather prominent rounded simple IV 23 and V2 weakly developed inner metatarsaltubercle distinct (IMT 09) oval more than2 times in length of toe I (ITL 19) supernu-merary tubercle present on all toes (fig 3E)Skin of snout and between eyes shagreened up-per eyelids slightly granular side of head espe-cially tympanic region granular (fig 3C) backand flanks granular dorsal part of forelimbsshagreened to sparsely granular thigh and tibiagranular tarsus shagreened ventral part of fore-limbs granular throat shagreened to granularchest belly and posterior surface of thighs gran-ular (fig 3B)

Colour in life Dorsally pale grayish greentinged with shining silver and cobalt violet adark grey inverted lsquoVrsquo on the dorsum runningfrom the level of the eye to nearly the level

of the vent (fig 3A) with black brown andsilver violet spots of variable size present es-pecially on flanks loreal and temporal regionbrownish black iris golden brown with a green-ish brown outer ring pupil with a golden outerring limbs light grey-brown with blackish crossbands finger tips white femur coffee brownwith variable-sized light brown and yellowishbrown blotches tibia and tarsus dark brownwith grayish cross bands toe discs white ven-trally with variable amounts of dark and lightpatches and spots side of the belly with lightyellowish brown blotches ventral surface ofthigh tibia and tarsus grayish brown intermin-gled with brown blotches feet and hands darkgrayish brown with black spots

Colour in alcohol Dorsum light grayishbrown black and grey spots throughout in-

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

8 SD Biju F Bossuyt

verted lsquoVrsquo dark grey loreal and tympanic regiondark brown forelimbs and hind limbs with lightgrey and light brown cross bands femur lightbrown with light grey blotches tibia and tarsuslight brown ventrally uniform grayish brownin different intensity throat light brown withminute black spots lateral side of the abdomenwith grayish white blotches surrounded by darkblackish brown area ventral surface of thightibia and tarsus brown intermingled with blackspots feet and hands light grayish brown withblack spots

Secondary male sexual characters Mediansubgular vocal sac present a pair of openingspresent at the base of lower jaw

Secondary female sexual characters Oviductof both females zigzag translucent ovary largecreamy white eggs about 3 mm in diameter

Etymology This species is named after AlainDubois of the Paris Museum of Natural HistoryFrance as a token of appreciation for his con-tribution in batrachological research in generaland his guidance to SDB and FB in their firststeps in batrachology in particular The speciesname is used as a noun in the nominative singu-lar standing in apposition to the generic name

Variation Measurements (in mm) of the typeseries are given in table 1 These frogs arehighly variable in coloration dorsally grey-white light green yellowish green dark cobaltgreen dark orange reddish brown reddishbrown or brownish green in all the colourforms one inverted lsquoVrsquo or two concave lines orscattered spots extend from the posterior borderof orbit to the vent BNHS 4286 uniform lightbrown dorsum with continuous faint concavemarking (fig 4A) BNHS 4287 light browndorsum and grey lateral side with prominentdark granular projections throughout (fig 4B)BNHS 4284 dark red-brown with black gran-ular projections in the shape of a concave linefrom behind the eye to the level of the vent faintgrey connection between eyes BNHS 4282light greenish-yellow discontinuous concave

line and faint grey connection between eyeBNHS 4285 grey-white with faint concave lineand a grey connection between eye The Limbcolour varies in a similar way as the dorsalcolour the colour during the breeding period isgenerally brighter than that of the non-breedingperiod (hibernating under stones or crevices)Palmar and supernumerary tubercles on fingersare of different size and prominent in severalspecimens The skin texture also shows con-siderable variation especially in female speci-mens One female (BNHS 4286) collected fromrock crevices has less granular projections onflanks (fig 4A)

Distribution and Natural History Philautusdubois is currently known only from the type lo-cality in Kodaikanal south of the lsquoPalghat GaprsquoThe holotype and six paratypes were collectedduring a rainy night from leaves about one me-ter high in shrubs near the roadside

Discussion

Although frogs of the genus Philautus can befound in a wide range of habitats the high-est species densities in Peninsular India arein the rainforests of the Western Ghats (Biju2001) where climatic conditions for their di-rect development are ideal Because this genusis a speciose group of small frogs with some-times highly variable coloration (Inger et al1984 Bossuyt and Dubois 2001) it has beenconsidered a particularly difficult group fortaxonomists We undertook an extensive sur-vey throughout the Western Ghats for the lastdecade and documented not only the distrib-ution of known species but also found manyundescribed taxa (Biju 2001) Museum stud-ies further demonstrated that at least fifty per-cent of the undescribed species we discoveredwere in fact already present in collections un-der an incorrect name In addition our stud-ies revealed that a large part of the litera-ture on identification and distribution of In-dian Philautus is highly questionable The two

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)

Two new species of Philautus 9

new species described herein perfectly fit in theabove described framework Indeed both Phi-lautus anili and P dubois were collected fromthe township of Wayanad and Kodaikanal re-spectively where they are abundant not onlyin rainforests but also in microhabitats rangingfrom roadside vegetation to gardens This illus-trates that indeed several of the most commonWestern Ghats anuran species are not scientifi-cally recorded yet and that there is still a longway to go before the richness of the WesternGhats fauna will be fully described

Acknowledgements The authors are very grateful toJRB Alfred and MS Ravichandran (ZSI) B Clarke andM Wilkinson (BMNH) Asad Rehmani and Varad Giri(BNHS) A Dubois and A Ohler (MNHN) H Grillitschand F Tiedemann (NMW) R Guumlnther (ZMB) RobertF Inger HK Voris and A Resetar (FMNH) and RohanPethiyagoda (WHT) for giving access to specimen in theircollections SDB is grateful to the Museacuteum National drsquo His-toire Naturelle Paris (MNHN) fellowship 2001 Conser-vation International (CI) Washington USA the Indian Na-tional Science Academy (INSA) and the Royal Society Lon-don fellowship 2003 for travel and museum studies RobertF Inger gave critical comments on an earlier version ofthe manuscript The Kerala Forest Department and TamilNadu Forest Departments kindly gave collection and studypermits Franky Bossuyt is a postdoctoral researcher at theFonds voor Wetenschappelijk Onderzoek Vlaanderen

References

Biju SD (2001) A synopsis to the frog fauna of theWestern Ghats India Indian Soc Conserv Biol OccasPubl 1 1-24

Biju SD Bossuyt F (2003) New frog family from In-dia reveals an ancient biogeographical link with the Sey-chelles Nature 425 711-714

Bossuyt F Dubois A (2001) A review of the frog genusPhilautus Gistel 1848 (Amphibia Anura RanidaeRhacophorinae) Zeylanica 6 1-112

Bossuyt F (2002) A new species of Philautus (AnuraRanidae) from the Western Ghats of India J Herpetol36 656-661

Das I (2000) Updates in amphibian systematics andnomenclature for the Indian region Biodiversity India8-12 3-4

Dutta SK (1997) Amphibians of India and Sri Lanka(Checklist and bibliography) Odyssey PublishingHouse Bhubaneswar India

Frost DF (2004) Amphibian species of the worldan online reference V30 httpresearchamnhorgherpetologyamphibiaindexhtml (American Museumof Natural History 22 August 2004)

Inger RF Shaffer HB Koshy M Bakde R (1984) Areport on a collection of amphibians and reptiles fromthe Ponmudi Kerala South India J Bombay nat HistSoc 81 551-570

Inger RF (1999) Distribution of amphibians in SouthernAsia and adjacent islands In Patterns of Distributionof Amphibia p 445-482 Duellman WE Ed JohnsHopkins University press Baltimore

Krishnamurthy SV Manjunatha Reddy AH GururajaKV (2001) A new species of frog in the genus Nyctiba-trachus (Anura Ranidae) from Western Ghats IndiaCurr Sci 80 887-891

Kuramoto M Joshy SH (2003) Two new species ofthe genus Philautus (Anura Rhacophoridae) from theWestern Ghats Southwestern India Curr Herpetol 2251-60

Ravichandran MS (1997) A new frog of the genusNyctibatrachus (Anura Ranidae) from southern IndiaHamadryad 22 9-12

Roelants K Jiang J Bossuyt F (2004) Endemic ranid(Amphibia Anura) genera in southern mountain rangesof the Indian subcontinent represents ancient frog lin-eage evidence from molecular data Mol PhylogenetEvol 31 730-740

Received July 6 2004 Accepted November 9 2004

Appendix 1

Specimens Examined

Philautus annandalii (BMNH 194722658) P beddomii(BMNH 194722659-66 NMW 22884 SDB 456) P bom-bayensis (ZSIC 18287 SDB 40175) P chalazodes (BMNH19472635) P charius (MNHN 19995597 SDB 4210-13)P flaviventris (BMNH 194722698) P garo (ZSIC 19187)P glandulosus (BMNH 194722722 SDB 40239) P griet(KBIN 1919-1922 1926 SDB 465) P jerdonii (BMNH19472784) P kempiae (BMNH collection number un-known) P luteolus (BNHS 4191 SDB 1126-1128) P mi-crodiscus (ZSIC 16924) P namdaphaensis (ZSIC A7177)P sanctisilvaticus (ZSIC A1778) P lsquoshillongensisrsquo (ZSICA6971 SDB 1280) P signatus (BMNH 194722736SDB 4006) P terebrans (USNM 239428) P similipalen-sis (ZSI A 9061) P tinniens (MNHN 19850527 SDB523-524) P travancoricus (BMNH 19472620 SDB4500) P tuberohumerus (BNHS 4193-94 SDB1101-1104)P wynaadensis (MNHN 1999 5596 SDB 3010-11)