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A New Species of Heterixalus (Amphibia: Hyperoliidae) from Western Madagascar

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A New Species of Heterixalus (Amphibia: Hyperoliidae) from Western Madagascar A new species of Heterixalus (Amphibia: Hyperoliidae) from western Madagascar Miguel Vences1*, Frank Glaw2, Riccardo Jesu3 & Giovanni Schimmenti3 1Zoologisches Forschungsinstitut und Museum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany E-mail: [email protected] 2Zoologische Staatssammlung, Münchhausenstr. 21, 81247 München, Germany 3Acquario di Genova, Area Porto Antico, Ponte Spinola, 16128 Genova, Italy Received 10 January 2000. Accepted 26 April 2000 A new species of hyperoliid frog, Heterixalus carbonei n.sp., is described from the Antsingy forest in western Madagascar. It is characterized by an advertisement call consisting of long and regular note series. The only other Heterixalus with a similar call structure, H. betsileo from higher altitudes in eastern Madagascar, has distinctly shorter notes. After recent discov- eries of the occurrence of H. luteostriatus and H. tricolor in western Madagascar, the new spe- cies brings the number of Heterixalus known from western Madagascar to three. Key words: Amphibia: Hyperoliidae: Heterixalus carbonei n.sp., Tsingy de Bemaraha, biogeography. INTRODUCTION new species of reptiles, some of which may be en- Frogs of the family Hyperoliidae are currently demic to the area. classified in 19 genera. Seventeen of these are en- demic to sub-Saharan Africa (Schirtz 1999). In ad- STUDY AREA dition, the monotypic genus Tachycnemis occurs on Situated in the centre of western Madagascar the Seychelles Islands, and the genus Heterixalus (18°15’–19°20’S; 44°15’–45°05’E), the World Heritage (currently 10 species) is endemic to Madagascar Site ‘Tsingy de Bemaraha’ consists of 500 000 ha, (Drewes 1984; Richards & Moore 1996). most of which are covered with secondary vegeta- Except for the highland species H. rutenbergi, all tion. The Plateau of Bemaraha (covered by the pri- Heterixalus are uniform in morphology (Glaw & mary deciduous Antsingy forest at its western Vences 1994). Colour patterns, on the other hand, slopes) as well as the grasslands, the patches of are variable within and among populations of forest and the mangroves situated between the single Heterixalus species (Glaw & Vences 1993, plateau and the Mozambique Channel, are all 1994). This situation is also typical for other included in this poorly known area. The limits hyperoliids (Schirtz 1999). On the other hand, of the Strict Nature Reserve N 9 ‘Tsingy de many Heterixalus species are bioacoustically well Bemaraha’, established in 1927, coincide almost differentiated (except the closely related and pos- perfectly with the ones of the Antsingy forest. sibly conspecific H. boettgeri, H. alboguttatus, and H. The climate is characterized by a marked contrast madagascariensis, as well as H. tricolor, H. variabilis, between the warm dry season (from May to Octo- and H. andrakata; Blommers-Schlösser 1982; Glaw ber) and the hot rainy season (from November to & Vences 1991, 1993, 1994; Andreone et al. 1994); April). In the Antsingy forest, which represents the advertisement calls are therefore the most useful wetter part of the area, annual rainfall is 1000– tool to distinguish different species of the genus in 1500 mm and the mean annual temperature the field. ranges between 25 and 28 °C (Bousquet & Rabe- In the present paper a new species of Heterixalus taliana 1992). is described from the ‘Tsingy de Bemaraha’ massif The eastern territories of the area, which are part in western Madagascar. In recent years several of the sedimentary region of Madagascar, are surveys (Emanueli & Jesu 1995; Schimmenti & mainly formed by a massif (Tsingy de Bemaraha) Jesu 1996, 1997; Jesu et al. 1999) of the unique habi- consisting of Upper Jurassic limestone. This massif tats within this massif have led to the discovery of extends north–south for more than 80 km with an *Author for correspondence. average width of 20 km and a maximum altitude of 35(2): 269–276 (October 2000) 270 African Zoology Vol. 35, No. 2, October 2000 934 m above sea level (a.s.l.). The dissected relief is Heterixalus carbonei n.sp., Figs 1–5 markedly karstic and shows typical erosion shapes consisting of limestone blocks with sharp Etymology. Dedicated to the memory of Ermanno tips at the top (locally named ‘tsingy’) and exten- Carbone, last Honorary Consul of Madagascar in sive cave systems. Genova, this new tree frog species will always The abundance of springs and streams allows remind us of his enthusiastic willingness to help the presence of a flourishing vegetation, which and his endless pleasantness. has never been affected by deforestation due Diagnosis. A species of the genus Heterixalus as to the impenetrability of the karstic habitat indicated by a vertical pupil, presence of a gular (Bousquet & Rabetaliana 1992). The Antsingy for- gland, connected lateral metatarsalia, and absence est, which represents at least5%oftotal western of spinulous skin in males which is typical in Madagascar deciduous forest cover, shows an Afrixalus (Drewes 1984). Distinguished from all extraordinary variety of facies in relation to substra- Heterixalus except H. betsileo by the general struc- tum nature, hydrography and exposition. Exten- ture of the advertisement call which consists of a sive descriptions of these habitats are given by single unharmonious note type repeated after Bousquet & Rabetaliana (1992) and Emanueli & regular intervals. Distinguished from H. betsileo by Jesu (1995). the distinctly longer note duration of advertise- ment calls. Further distinguished from H. ruten- MATERIALS & METHODS bergi by the lack of black pigment on the gular Fieldwork was carried out by R.J. and G.S. during gland and lack of a medial dorsal stripe; from the rainy season (25–29 January 1995 and 2–14 H. madagascariensis, H. boettgeri, H. alboguttatus, March 1997) at two sites along the western slopes and H. punctatus by the presence of light dorso- of the Antsingy forest: Trano Passage (18°43’S, lateral bands in adults; and from H. andrakata by 47°07’E; 70 m a.s.l.) and Ambodyreana (18°41’S, the lack of dark dorsal marbling and vermicula- 47°07’E; 120–150 m a.s.l.). The specimens of the tions. new species described in this paper were observed Holotype. Adult male (MSNG 49090; Figs 1, 2), and collected only at the former site. The search collected 13 March 1997 at Trano Passage, Strict for frogs was carried out after sunset when males Nature Reserve N 9 ‘Tsingy de Bemaraha’, 18°42’S, could be easily located while emitting their 44°43’E, 70 m a.s.l., Antsalova Fivondronana, advertisement calls, which were recorded using a Mahajanga Province, Madagascar, by Riccardo SONY TCM-77V tape recorder with external Jesu, Giovanni Schimmenti, Herinjato M. Rakoton- microphone; at the same time air temperature and drasoa, Paolo Bistagnino and Giacomo Berichilli. relative humidity were recorded. After exposure Other types. None. to acetic ether, the sacrificed specimens were fixed Description of the holotype. Body slender; head and preserved in 70 % ethanol. wider than long, slightly broader than body; snout Calls were analysed with the MEDAV sound rounded in dorsal and lateral views; nostrils analysing system Spektro 3.2. Morphological directed laterally, not protuberant; canthus ros- measurements were taken by the same person tralis indistinct, concave; loreal region more or less (M.V.)with a calliper to the nearest 0.1 mm. Speci- even; tympanum indistinct, rounded, very close mens were deposited in the Museo di Storia to eye; its diameter almost half of eye diameter; Naturale di Genova (MSNG). Webbing formula is supratympanic fold absent; tongue ovoid, dis- given according to Blommers-Schlösser (1979). tinctly bifid posteriorly; vomerine teeth absent, To ascertain the genetic differentiation of the choanae small, rounded; maxillary teeth present. new species to other Heterixalus, we sequenced a Arms slender; subarticular tubercles single; no fragment (555 bp) of the mitochondrial 16S rRNA visible outer or inner metacarpal tubercles; com- gene homologous to the base pair positions parative finger length 1<2<4<3; well-developed 4006–4543 of the Xenopus laevis mt genome (Roe terminal finger disks; webbing formula of hand 1 et al. 1985), using the primers and cycling protocols (1.5), 2i (2), 2e(1), 3i (2), 3e (2), 4 (1.5). Legs slender; given in Vences et al. (2000). Sequences were sub- tibiotarsal articulation reaches anterior corner of mitted to Genbank (accession numbers AF215433– eye when the limbs are adpressed along body; AF215437 and AF215502–AF215503). More de- feet with a small elliptical inner metatarsal tuber- tailed information on the genetic analyses will be cle and without recognizable outer metatarsal published in a forthcoming paper. tubercle; subarticular tubercles single; toe disks Vences et al.: New species of Heterixalus 271 Fig. 1. Holotype of Heterixalus carbonei n.sp. (MSNG 49090) from Trano Passage, Antsingy forest (diurnal markings). moderately expanded; webbing formula of foot 1 (Fig. 1). A single yellow-brown dorsolateral band (0.5), 2i(1), 2e(0.5), 3i(1), 3e(0.5), 4i(1), 4e(1.5), 5(0.5). on each side. Shanks and foot also partly yellow- Lateral metatarsalia connected; comparative toe brown. Inguinal region and lateral parts of limbs length 1<2<3<5<4. Finger and toe pads with yellow. Venter white, slightly translucent. Throat continuous circummarginal groove. Skin on the yellow, gular gland less brightly coloured than upper surface smooth; ventral surface of legs and surrounding portion of throat. Fingers and toes chest smooth, venter slightly granular. A large, ventrally yellow, webbing yellowish. In preserva- ovoid gular gland on throat, wider than long tive, back and dorsal surface of legs brown. (width 6.0 mm, length c. 4.7 mm), not heart- Dorsolateral stripes faintly recognizable as light shaped, without clear posterior border. First finger brown areas.Ventrally uniformly whitish. with a distinct whitish nuptial pad. Measurements of holotype. Snout-vent length Colour in life (at night) dorsally dirty brownish 26.4 mm; maximum head width 8.5 mm; head Fig.
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