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Zootaxa, Description of Tadpoles of Five Frog Species In TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Zootaxa 1988: 48–60 (2009) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition) Description of tadpoles of five frog species in the subgenus Brygoomantis from Madagascar (Mantellidae: Mantidactylus) HEIKE SCHMIDT1, AXEL STRAUß1, FRANK GLAW2, MEIKE TESCHKE3 & MIGUEL VENCES1,4 1Zoological Institute, Technical University of Braunschweig, Spielmannstr. 8, 38106 Braunschweig, Germany 2Zoologische Staatssammlung München, Münchhausenstr. 21, 81247 München, Germany 3Max-Planck Institute for Evolutionary Biology, Department of Evolutionary Genetics, 24306 Plön, Germany 4Corresponding author. E-mail [email protected] Abstract We describe the larval stages of five frog species classified in the Madagascan subgenus Brygoomantis of the genus Mantidactylus, which were identified by DNA barcoding: Mantidactylus alutus, Mantidactylus curtus, and three taxonomically undescribed species here named Mantidactylus sp. aff. biporus "Ranomafana", M. sp. aff. biporus "Marojejy", and M. sp. aff. curtus "Ankaratra". The larvae of M. alutus, and of M. sp. aff. curtus "Ankaratra", had been described before, and we confirm and complement the previous studies. Our data confirm that Brygoomantis tadpoles are benthic, of a rather generalized body shape and oral disc morphology. All species for which tadpoles are known so far have a large dorsal gap of marginal papillae, a labial tooth row formula of 3-5 rows on the anterior labium of which only the first is continuous and the others have distinct medial gaps, and three rows of labial teeth on the posterior labium of which the first usually has a very small medial gap which however can sometimes be undetectable. Total number of marginal and submarginal papillae ranges from 50 to 103 and density of labial teeth (on the second upper row) from 19 to 76 per mm. Key words: Amphibia, Anura, Mantidactylus alutus, Mantidactylus curtus, Mantidactylus sp. aff. biporus, Mantidactylus sp. aff. curtus, tadpole descriptions, DNA barcoding Introduction Madagascar harbours a fascinating variety of anuran amphibians that are still incompletely inventoried. Even less complete is the inventory of their larval forms, although these may be relevant to understand the adaptations in the endemic radiations of Malagasy frogs (e.g., Randrianiaina et al. 2007). Within the family Mantellidae, endemic to Madagascar, Mantidactylus Boulenger is one of the most species-rich lineages. The genus is subdivided into five subgenera (Glaw & Vences 2006). Species in one of these subgenera, Brygoomantis Dubois, were formerly known as the Mantidactylus ulcerous species group (Blommers- Schlösser 1979; Blommers-Schlösser & Blanc 1991). Brygoomantis are small to medium-sized frogs of usually brownish dorsal colour and inconspicuous appearance. These frogs are characterised by riparian to semiaquatic habits, i.e., they are usually found along streams and many of them typically escape by diving into the water. They can be found throughout Madagascar along streams, occasionally occurring also in lentic waters like swamps, displaying diurnal and nocturnal activity (Blommers-Schlösser 1979; Blommers- Schlösser & Blanc 1991; Glaw & Vences 1994, 2007). Some larvae assigned to species of Brygoomantis have been described by Arnoult & Razarihelisoa (1967) and Blommers-Schlösser (1979). However, due to the large number of cryptic, yet undescribed species in this subgenus (e.g., Glaw & Vences 2007), the exact assignment of the described tadpoles to species is in some cases questionable. In addition, some of these previous descriptions were based on series from different 48 Accepted by S.Carranza: 8 Dec. 2008; published: 27 Jan. 2009 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. localities which may represent more than one species. Recently, Knoll et al. (2007) provided data on the tadpoles of two described and two undescribed Brygoomantis species, based on specimens identified and matched to adults by DNA barcoding. As a contribution to the goal of a complete and comparable set of larval descriptions of Malagasy frogs (e.g., Thomas et al. 2005; Raharivololoniaina et al. 2006; Vejarano et al. 2006; Randrianiaina et al. 2007; Knoll et al. 2007) we here present descriptions of the previously unknown tadpoles of three species, and re- descriptions of two additional species of Brygoomantis. Materials and methods Tadpoles were collected in the field, euthanised by immersion in chlorobutanol solution, and immediately sorted into series based on their morphology. From each series one specimen was selected and a tissue sample from its tail musculature or fin taken and preserved in 99% ethanol. These specimens are here named “DNA vouchers. Drawings and descriptions of the tadpoles in this paper are based only on the DNA vouchers. All specimens of each series were examined to assess morphological variability and to complete the structures missing because of tissue sampling in the DNA vouchers. After tissue collection, specimens were preserved in 4% formalin, or in 70% ethanol in the case of the series ZSM 123/2007. Specimens were deposited in the Zoologische Staatssammlung München, Germany (ZSM). We give the ZSM catalogue number of voucher specimens as well as the field numbers (ZCMV; FGZC; FG/MV) applied to the same series. Tadpoles were identified using a DNA barcoding approach based on a fragment of the mitochondrial 16S rRNA gene, which is known to be sufficiently variable among species of Malagasy frogs (Thomas et al. 2005). The circa 550 bp were amplified using primers 16Sar-L and 16Sbr-H from Palumbi et al. (1991) applying standard protocols, resolved on automated sequencers, and compared to a near-complete database of sequences of adult Malagasy frog species. Identification was considered to be unequivocal when the tadpole sequence was 99-100% identical to an adult specimen from the same geographical region, and not more similar to any sequence from another species. DNA sequences were deposited in Genbank (accession numbers EU717879-EU717893; accession numbers of comparative adult specimens are included in the sequence sets AY847959-AY848683 and AJ315909-AJ315913). Developmental stages are based on Gosner (1960). Morphological terminology follows Altig & McDiarmid (1999). Labial tooth row formulae are given according to Altig (1970) and Altig & McDiarmid (1999). A1-A5 is used to refer to the anterior (upper) labial tooth rows, P1-P3 for the posterior (lower) rows. Small and irregularly occurring gaps in rows of labial teeth were interpreted as oral deformities (see Drake et al. 2007). Measurements of body and tail were taken using a digital calliper (effective range: 0-150 mm; precision: 0.03 mm; repeatability: 0.1 mm). Drawings are mainly based on the preserved tadpoles. Pictures and measurements of the oral apparatus were taken using a Zeiss Stereo Discovery microscope with a mounted digital camera. Abbreviations used in the descriptions are as follows: BL, body length (distance from the tip of the snout to the body-tail junction); BH, maximum body height; BW, maximum body width (at the widest point); TL, total length (sum of body length and tail length); TAL, tail length; ED, eye diameter; IOD, inter-orbital distance (measured between the centres of the pupils); IND, inter-narial distance (measured between the centres of the narial apertures); ESD, distance between eye and spiraculum (distance between the centre of the eye and the centre of the spiracular apertures); LTRF, labial tooth row formula; LTD, labial tooth density (number of teeth per millimetre in A2); LTN, number of all teeth in A2; PN, papillae number; TMH, tail muscle height (measured vertically from the junction of the body wall with the ventral margin of the tail muscle); TMW, tail muscle width (measured horizontally from the sinistral and the dextral junction of the body); MTH, maximum tail height (including fins and caudal musculature, taken at its maximal vertical extent); DF, height of dorsal fin at midlength of tail; TMHM, height of tail muscle at midlength of tail; VF, height of ventral fin at midlength of tail; ODW, oral disc width (maximum width of the oral disc). TADPOLES OF MANTIDACTYLUS (BRYGOOMANTIS) Zootaxa 1988 © 2009 Magnolia Press · 49 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Results Mantidactylus sp. aff. biporus ”Ranomafana Series examined. ZSM 123/2007. The 14 tadpoles of this series were collected in a rather shallow stream at a site locally known as Fompohonina in the Ranomafana National Park in eastern Madagascar (stream called Fompohonina III; depth about 7 cm, 21°15.907'S; 47°25.349'E, 1012 m a.s.l.) by A. Strauß, R.-D. Randrianiaina and S. H. Ndriantsoa on 16 March 2007. The stream had a low downward slope of 16 cm per 10 m and its substrate was characterised by a high percentage of sand (58%) and leaves (40%), almost without vegetation. It was surrounded by primary rain forest with high canopy cover (> 70%). Water temperature was 18.9°C, pH was 8.9. The tadpoles were collected within or close to leaf agglomerations in the slower running parts of the stream. Taxonomic note. The DNA sequence of the DNA voucher of
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