Raorchestes primarrumpfi, Vijayakumar, S. P., Dinesh, K. P., Prabhu, Mrugank V. & Shanker, Kartik, 2014

Vijayakumar, S. P., Dinesh, K. P., Prabhu, Mrugank V. & Shanker, Kartik, 2014, Lineage delimitation and description of nine new species of bush frogs (Anura: Raorchestes, Rhacophoridae) from the Western Ghats Escarpment, Zootaxa 3893 (4), pp. 451-488 : 479-484

9. Raorchestes primarrumpfi sp. nov.

( Figures 2 , 3 , 13 , 14 & 15 ; Tables 2 & 3)

Holotype: ZSI/ WGRC /V/A/881 ( CESF 1276), a male (SVL 21.0 mm), collected by S.P. Vijayakumar and Mayavan June 2011 from a grassland site (11.2331 N, 76.5443 E), Nilgiri Massif ( Fig 1 ), Western Ghats, Peninsular India.

Paratype: ZSI/ WGRC /V/A/882 ( CESF 441), a male (SVL 19.9 mm), collected by S.P. Vijayakumar, August 2009 from a grassland site (11.2331 N, 76.5443 E), Nilgiri Massif ( Fig 1 ), Western Ghats, Peninsular India.

Lineage diagnosis. Raorchestes primarrumpfi sp. nov. can be phylogenetically diagnosed as belonging to the Tinniens clade ( Fig 3 ), showing a well supported sister relationship with an another unidentified lineage occurring in sympatry. Despite its shallow genetic divergence (1–2 % on 16S gene) from its sympatric sister lineage, it exhibits high divergence in morphological characteristics both in the multivariate morphological space ( Fig 14 ) as well in the dorsal and ventral coloration ( Fig 13 ). Iris coloration and patterns ( Fig 13 (b)) were also found to show distinct differences from its sister lineage.

Field diagnosis. Raorchestes primarrumpfi sp. nov. can be distinguished from the related congeners by the following combination of characters.

Morphology. Raorchestes primarrumpfi sp. nov. can be distinguished by its (1) shorter tibia length (ShL/ SVL=0.32 (0.31–0.35, n=5) (vs. ShL/SVL=0.45 (0.43–46, n=3) in the unidentified lineage and ShL/SVL=0.41 (0.41–42, n=3) in R. tinniens ); (2) shorter thigh length (TL/SVL = 0.35, n=5) (vs. TL/SVL=0.45, n= 3 in the unidentified lineage and TL/SVL=0.44, n= 3 in R. tinniens ); (3) smaller size (males) (SVL=20 (18.4–21.0, n=5) (vs. 22 (21–22.8, n=3) in the unidentified lineage); (4) shorter head length (HL/SVL=0.26 (0.24–0.28, n=5) (vs. 0.37 (0.35–0.38, n=3) in R. tinniens ; (5) shorter snout length (SL/SVL=0.11 (0.10–0.13, n=5) (vs. 0.15, n=3) in the unidentified lineage); (6) unique iris coloration, lower part dark maroon and upper half speckled with iridescent golden and silvery colour (vs. uniform brown in the unidentified lineage and uniform brown with golden speckles in R. tinniens ; (7) dorsum largely granular, dark olive and with a consistent pattern of three distinct maroon longitudinal discontinuous stripes (vs. highly variable dorsum coloration with no distinct pattern in the related lineages); (8) two distinct maroon blotches on the eyelids extending slightly into inter-orbital space (vs. absent in the other lineages) (9) ventral coloration is white with a bluish wash towards sides (vs. shades of yellow, areolate skin, semi-transparent with yellow blotches in some individuals and shades of yellow with black spots in R. tinniens (Biju and Bossuyt 2009)) .

Geography. Restricted in range to the very high elevations of the Nilgiri Massif (see natural history and distribution for details). Overlaps broadly in its geographical range with an unidentified lineage and R. signatus . Ecology. Observed to be restricted to grasslands and swamps.

Description of holotype (all measurements in mm). A small sized squat bush frog (SVL = 21.0 mm), width of head broader than head length (HW = 7.5 mm; HL = 5.5 mm), arched, flat dorsally; snout rounded in total profile, slightly protruding beyond mouth. Snout length is equal to diameter of eye (SL = 2.5 mm, EL = 2.4 mm). Canthus rostralis rounded, loreal region slightly concave. Interorbital space (IUE = 2.2 mm) flat and sub equal to upper eyelid (UEW = 1.8 mm). Interorbital space between posterior margins of the eyes 1.9 times that of anterior margins (IFE = 3.5, IBE = 6.6 mm). Nostrils oval, nearer to tip of snout. Weak symphysial knob. Pupil horizontal. Tympanum indistinct. Tongue bifid, granular with a papilla. Supratympanic fold from behind eye to shoulder.

Relative length of fingers I<II<IV<III. Finger tips with small disks (fd3 = 0.6 mm, fw3 = 0.6 mm) with distinct circum–marginal grooves, fingers with dermal fringes on both sides. Webbing on palm absent, subarticular tubercles indistinct, pre-pollex indistinct and supernumerary tubercles absent.

Hind limb short, heels fall apart when folded at right angles to the body. Thigh/Femur (TL = 7.0 mm), sub equal to Shank/Tibia (ShL = 6.5 mm) and foot (FOL = 6.6 mm) but less than heel to tip of fourth toe (TFOL = 10.8 mm). Relative toe length I<II<III<V<IV, webbing rudimentary, web formula (I 1- 1 II 1- 2 III 1½- 3 IV 2½-1½ V). Tibiotarsal articulation reaches shoulder region. Outer metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent.

Color in life. Dorsum, background olive with a dark maroon longitudinal disconnected striped pattern, the pattern extending on to outer two fingers on the forelimb and femur, tibia, tarsus and outer toe on the hind limb. Canthus ridge darker extending to the tip of snout and flesh coloured, with olive patches, in a few individuals. Laterally, behind shoulders a distinct flesh coloured patch hidden in resting position; bluish white small blotches along the lateral sides and distinct blotches towards groin and anterior femur on a maroon background, this pattern is variable across individuals. Ventrally white, with a light bluish wash towards lateral edges. Ventral parts of tibia and tarsus with elongated white patches on a fleshy background. Iris, lower 1/3rd dark maroon, upper half speckled with iridescent golden and silvery colour and outer posterior orbital ring blue.

Etymology. Derived and modified from ‘ Primarrumpf’, a German term used by geomorphologists to refer to remnant primitive surfaces of Gondwanaland. In the Western Ghats Escarpment, these surfaces occur in the Nilgiri and Anaimalai massifs.

Natural history and distribution. All the calling males were observed amidst dense grass clumps and herbs in the montane grasslands and the detection of this species was higher in swampy grasslands ( Fig 15 ). It exhibits a narrow geographical range and is restricted in distribution to the montane zone (2212−2359 m, n=13) towards the western edge of the Nilgiri Massif ( Fig 1 & 2 ). The higher elevations of the Camels Hump Massif, adjacent to the Nilgiri Massif, might hold a relative of this lineage and needs further exploration.

Sl. No Species

1 Raorchestes agasthyaensis Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 2 Raorchestes akroparallagi ( Biju and Bossuyt, 2009) 3 Raorchestes anili ( Biju and Bossuyt, 2006)

4 Raorchestes archaeos sp. nov.

5 Raorchestes aureus sp. nov.

6 Raorchestes beddomii (Gunther, 1876)

7 Raorchestes blandus sp. nov.

8 Raorchestes bobingeri (Biju and Bossuyt, 2005)

9 Raorchestes bombayensis ( Annandale, 1919)

10 Raorchestes chalazodes (Gunther, 1876)

11 Raorchestes charius ( Rao, 1937)

12 Raorchestes chlorosomma ( Biju and Bossuyt, 2009) 13 Raorchestes chotta ( Biju and Bossuyt, 2009)

14 Raorchestes chromasynchysi ( Biju and Bossuyt, 2009) 15 Raorchestes coonoorensis ( Biju and Bossuyt, 2009) 16 Raorchestes crustai Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 17 Raorchestes dubois ( Biju and Bossuyt, 2006)

18 Raorchestes echinatus sp. nov.

19 Raorchestes emeraldi sp. nov.

20 Raorchestes flaviventris (Boulenger, 1882) *

21 Raorchestes flaviocularis sp. nov.

22 Raorchestes ghatei Padhye, Sayyed, Jadhav and Dahanukar, 2013 23 Raorchestes glandulosus (Jerdon, 1853)

24 Raorchestes graminirupes (Biju and Bossuyt, 2005) 25 Raorchestes griet (Bossuyt, 2002)

26 Raorchestes hassanensis ( Rao, 1937) #

27 Raorchestes indigo sp. nov.

28 Raorchestes jayarami ( Biju and Bossuyt, 2009)

29 Raorchestes johnceei Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 30 Raorchestes kadalarensis Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 31 Raorchestes kaikatti ( Biju and Bossuyt, 2009)

32 Raorchestes kakachi Seshadri, Gururaja and Aravind, 2012 33 Raorchestes leucolatus sp. nov.

34 Raorchestes luteolus ( Kuramoto and Joshy, 2003)

35 Raorchestes manohari Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 36 Raorchestes marki ( Biju and Bossuyt, 2009)

37 Raorchestes montanus (Jerdon, 1875) #

......continued on the next page Sl. No Species

38 Raorchestes munnarensis ( Biju and Bossuyt, 2009)

39 Raorchestes nerostagona (Biju and Bossuyt, 2005)

40 Raorchestes ochlandrae ( Gururaja, Dinesh, Palot, Radhakrishnan and Ramachandra, 2007) 41 Raorchestes ponmudi (Biju and Bossuyt, 2005)

42 Raorchestes primarrumpfi sp. nov.

43 Raorchestes ravii Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 44 Raorchestes resplendens Biju, Shouche, Dubois, Dutta and Bossuyt, 2010 45 Raorchestes signatus (Boulenger, 1882)

46 Raorchestes sushili ( Biju and Bossuyt, 2009)

47 Raorchestes theuerkaufi Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 48 Raorchestes thodai Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011 * 49 Raorchestes tinniens (Jerdon, 1853)

50 Raorchestes travancoricus ( Boulenger, 1891)

51 Raorchestes tuberohumerus ( Kuramoto and Joshy, 2003)

52 Raorchestes uthamani Zachariah, Dinesh, Kunhikrishnan, Das, Raju, Radhakrishnan, Palot and Kalesh, 2011

* species not included in the phylogenetic tree; # details of revalidation will be dealt elsewhere (under preparation)

Figures

FIGURE 2. Geographic distribution of new species of bush frogs (Raorchestes) from Western Ghats. Orange filled circles

FIGURE 13. Raorchestes primarrumpfi sp. nov. (a) profile in life; (b) eye profile in life; (c) dorsal profile of holotype; (d) ventral profile (in life); (e) R. UI 427 (life); (f) R. tinniens (in life); (g) R. UI 427 (eye profile—in life); (h) R. UI 427 (ventral profile—in life)

FIGURE 14. Separation of male individuals of Raorchestes primarrumpfi sp. nov. (triangle) and Raorchestes UI 427 (circle) in multivariate space. Axes represent principal component scores.

FIGURE 3. A maximum likelihood tree for bush frogs, and in the Western Ghats, based on three MtDNA genes: 16 S, Cytb and ND 1. The lineages described are marked in red. Sub-clades are named on the right.

FIGURE 15. A typical habitat of Raorchestes primarrumpfi sp. nov. Detection of the species was greater in the swampy grasslands. Locality: Avalanche zone, Nilgiri Massif.

FIGURE 1. A map of the Western Ghats showing the major physiographic features referred to in the text.