Mantidactylus kathrinae, Glaw & Vences & Gossmann, 2000

Glaw, Frank, Vences, Miguel & Gossmann, Viola, 2000, A new species of Mantidactylus (subgenus Guibemantis) from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae), Journal of Natural History 34 (7), pp. 1135-1154 : 1136-1144

Mantidactylus kathrinae n. sp.

HOLOTYPE ZFMK 62264 (®gures 1±2), adult male, collected in rainforest near An’ Ala (18 ss 56 ¾ S, 48 ss 28 ¾ E, 840 m above sea level), eastern Madagascar, on 3 February 1996 by F. Glaw.

PARATYPES ZFMK 62263 , adult male, same locality, date and collector as holotype ; ZFMK 62266 , same locality as holotype, collected on 30 January 1996 by D. Vallan . Additional specimens. ZFMK 64543 and 64544 from Andapa (14 ss 40 ¾ S,

49 ss 39 ¾ E), north-eastern Madagascar, are attributed to the new species but not de®ned as paratypes due to their bad state of preservation .

Diagnosis. A species of the genus Mantidactylus as indicated by the lack of nuptial pads in breeding males and by the presence of femoral glands (recognizable from internal view). A member of the subgenus Guibemantis as indicated by large relative hand length (hand length/snout ±vent length 0.32±0.35), a white vocal sac, femoral gland structure ( table 2) and arboreal egg deposition site above stagnant water bodies. The new species is distinguished from all other valid species of the subgenus Guibemantis ( M. depressiceps , M. tornieri , M. liber ) by a more distinct prepollex and by advertisement calls (®gures 3±6). It further diOEers from M. tornieri and M. liber by white eggs and white jelly of its clutches, from M. liber and M. depressiceps by its larger snout±vent length (adult males> 44 mm SVL) and from M. depressiceps by a more extended foot webbing (reaching further than last subarticular tubercle on ®fth toe), lower pulse repetition rate, longer note duration and lower dominant frequency ( table 3). Specimens from a population sympatric (but not syntopic) with M. depressiceps and M. tornieri can be easily diagnosed by their much larger size (adult males> 55 mm SVL).

Description of holotype. Measurements of the holotype are included in table 1. Body relatively slender; head longer than wide, wider than body; snout rather pointed in dorsal and lateral views; nostrils directed laterally, not protuberant; canthus rostralis distinct, straight; loreal region concave; tympanum distinct, small, rounded, its diameter about one-half that of eye; distinct supratympani c fold, running straight for most of its length, curving towards the forelimb insertion; tongue ovoid, distinctly bi®d posteriorly; vomerine teeth form a distinct, large, oblique group posterolateral to choanae; choanae medium-sized, rounded. Forelimbs slender; subarticular tubercles single; inner and outer metacarpal tubercle present; a distinct prepollex (3 mm) of about one-third of the length of the inner ®nger. Fingers with very little rudiments of webbing; relative ®nger length 1 <2 <4 <3; ®nger discs strongly enlarged; nuptial pads absent; ventral circummarginal groove of ®ngers and toes complete, very distinct.

Legs slender; tibiotarsal articulation reaches between eye and nostril; lateral metatarsalia separated; inner metatarsal tubercle rather small; outer metatarsal tubercle present, distinct; webbing formula of the foot 1 (1), 2i(1.5±2), 2e(0.75), 3i(1.75), 3e (0.75), 4i(2), 4e (1.75), 5(0.5) (since only one subarticular tubercle is recognizable on the second toe, the actual relative extension of the web can only be estimated on this toe); relative toe length 1 <2 <3<5 <4. Skin on the dorsum smooth; ventral skin smooth, but more granular posteriorly. Skin ventrally on thigh distinctly granular (glandular patch only slightly prominent). The gland extends over about three-quarters of the surface of the thigh and gland patches on opposite thighs are in contact in the anal region. There are no enlarged tubercles around the anus.

Colour of holotype in life. Dorsal colour fading from a lighter brown posteriorly to dark brown anteriorly, with a few very small scattered whitish spots. Head sides and tympanic region dark brown. Iris uniformly dark except a narrow copper streak along the upper margin. Webbing, posterodorsal part of thighs, arm insertions and lower ¯anks dark grey. On the ¯anks, the beige dorsal colour fades gradually into the more whitish ventral colour, with an irregular series of rather small light spots which are indistinctly bordered by darker pigment. Venter dirty white with some dark pigment, throat bright white, sharply bordering on the dark brown lower lip. Ventral side of fore-and hindlimbs greyish except the more yellowish thigh patch area.

Colour in preservative similar to that in life. Dorsum almost uniformly light brown to beige, dark colour of webbing less distinct.

Variation. The male paratypes are very similar to the holotype. The pattern of light spots (bordered by dark brown) on the ¯anks and the posterodorsal surface of the thigh is more contrasting in ZFMK 62266 . Canthus rostralis and supratympani c fold are light beige in this specimen, emphasizing the contrast between dark head side and light dorsum. In life, this specimen had a lighter dorsal colour (back and upper head surface light brown). Nearly the upper third of the iris was light brown to copper. Webbing formula of the foot is 1(1), 2i(1.5±2), 2e (0), 3i (1.5), 3e (0.5), 4i(2), 4e(1.75), 5 (0.5) in ZFMK 62263 and 1(1), 2i(1.5±2), 2e (0.5), 3i (1.75), 3e (0.75), 4i(2), 4e (2), 5 (0.5) in ZFMK 62266 . Of the additional specimens from Andapa, ZFMK 64543 (male) measures 44.4 mm SVL and ZFMK 64544 (female) 46.1 mm SVL.

The similar SVL of both Andapa specimens of M. kathrinae is in accordance with Blommers-SchloÈsser’s (1979) observation that in the subgenus Guibemantis sexual size dimorphism is very low. Although the available Andapa specimens are distinctly smaller than those from An’Ala, they still are larger than the available specimens of M. depressiceps . According to Blommers-SchloÈsser (1979), only one out of 11 examined M. depressiceps males, ZMA 6976, reached a SVL of 44 mm, the remaining ten specimens ranging from 34±41 mm.

Femoral glands. The structure of the femoral glands was studied in one paratype ( ZFMK 62266). From internal view it is evident that the externally visible granular patch is composed of a very large number of single light small granules. In the centre of the gland, these granules are densely arranged next to each other, without further structure, whereas towards the edges of the gland patch they are increasingly arranged in small rosette-like groups (®ve to seven granules surrounding one central granule). In some of these rosette-like granule groups, the central granule is dark, giving the impression of a central porus-like structure. For measurements of the gland structures see table 2.

Advertisement call. Calls were recorded at An’Ala (on 3 February 1995, c. 21:00 h) and at Andapa (by K. Schmidt on 18 March 1997, 18:00 h). Calls from both localities were rather similar and consisted of series of three to four pulsed notes (®gures 3±4, table 3).

Natural history. About 20 whitish clutches, very probably laid by M. kathrinae , were seen on 3 February 1996 at An’Ala hanging from the vegetation above a pool at few metres distance from a forest brook. On this evening, however, very little calling activity of M. kathrinae was observed. K. Schmidt (personal communication) observed many whitish clutches being laid by a large breeding colony of the species near Andapa in 1996, but heard only weak calling and breeding activity at the same pool on 18 March 1997. The senior author observed white clutches on 30 March 1994 (with greenish tadpoles ready to hatch) and on 23 February 1995 above a pond near the edge of a river in the Marojezy Reserve (c. 300 m above sea level) in northeastern Madagascar; these clutches may also have belonged to M. kathrinae .

Distribution. The new species is so far only known from the type locality and from Andapa in north-eastern Madagascar.

Etymology. Dedicated to Kathrin Schmidt, Bonn, in recognition for her invaluable help with our research on Malagasy amphibians and reptiles; her call recordings from Andapa helped to clarify the relationships and distribution of the species of the M. depressiceps complex.

Available names in Guibemantis

Rhacophorus mocquardii Boulenger, 1896 and Mantidactylus acuticeps Ahl, 1929 were considered to be junior synonyms of M. depressiceps (Blommers-SchloÈsser and Blanc, 1991; Glaw and Vences, 1992b). A re-examination of the holotypes of both taxa con®rmed the validity of these synonymizations (see below). We also re-examined the holotype of Rhacophorus tornieri Ahl, 1929 and the lectotype of Rhacophorus depressiceps Boulenger, 1882 to con®rm that the names Mantidactylus tornieri and M. depressiceps were correctly applied and de®ned by Blommers- SchloÈsser (1979) who ®rst noted the presence of two distinct biological species previously subsumed under the name depressiceps .

Measurements of the respective type specimens ( tornieri : holotype ZMB 30533 (®gure 7); depressiceps : lectotype BMNH 1947.2.27.5 0 (®gure 8), paralectotypes BMNH 1947.2.27.51 ±53; acuticeps : holotype ZMB 30496 (®gure 9); mocquardii : holotype BMNH 1947.2.8.6 2 (®gure 10 )) are included in table 1. Webbing formulae of the name-bearing types (holotypes of tornieri , acuticeps and mocquardii lectotype of depressiceps ) are 1(1), 2i(1), 2e(0.5), 3i(1.75), 3e (1), 4i(2), 4e (2), 5 (0.5) in tornieri ; 1(1), 2i(1.75), 2e(1), 3i(2), 3e (1), 4i(2.25), 4e (2.25), 5(1) in depressiceps ; 1 (1), 2i(1.75), 2e (0.5), 3i(2), 3e(1), 4i(2.25), 4e (2.25), 5(1) in acuticeps ; 1(1), 2i(1.25), 2e(0.75), 3i(2), 3e (1), 4i(2.25), 4e (2), 5 (0.75±1) in mocquardii .

The type specimens of acuticeps and depressiceps show reduced webbing and a small SVL compared with tornieri and kathrinae , con®rming the attribution of acuticeps as junior synonym of depressiceps . The holotype of mocquardii (maybe an immature specimen) is in a rather bad state of preservation and has slightly more webbing than generally present in depressiceps . We here consider it to be a junior synonym of M. depressiceps since the extent of webbing in mocquardi is less than in tornieri and its small size is a further diOEerence to that species. All four name bearing type specimens ( tornieri , depressiceps , acuticeps , mocquardii ) have a rather small and indistinct prepollex diOEerentiating them from M. kathrinae .