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Description of two Werneria tadpoles from Cameroon (Amphibia: Anura: Bufonidae) MAREIKE HIRSCHFELD1, MICHAEL F. BAREJ1, SIMON P. LOADER2 & MARK-OLIVER RÖDEL1 1
Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstraße 43, 10115 Berlin, Germany 2 Institute of Biogeography, University of Basel, Basel, Switzerland. E-mails:
[email protected],
[email protected],
[email protected],
[email protected]
Detailed tadpole descriptions for toads of the genus Werneria Poche, 1903 have been only published for W. preussi (Matschie, 1893) (Mertens 1938) and W. tandyi (Amiet, 1972) (Rödel et al. 2004), the latter description assigned to this species with reservation. A few characters of W. bambutensis (Amiet, 1972) tadpoles were mentioned by Amiet (1972). So far the assignment of tadpoles to a particular Werneria species was based on geography (locality and altitude). Three Werneria species, W. mertensiana, W. tandyi and W. bambutensis (Fig. 1), co-occur on Mount Manengouba in south-western Cameroon. There they inhabit forests and river edges in different but slightly overlapping altitudes (Rödel et al. 2004). During recent field work (November / December 2010) on Mount Manengouba we collected Werneria tadpoles which are herein described.
FIGURE 1. Three Werneria species live at different altitudes on Mount Manengouba (d), Cameroon. W. mertenisana (a, ZMB 76693) occurs at lowest altitudes (950–1350 m asl), followed by W. tandyi (b, ZMB 76697; 1200–1750 m asl) and W. bambutensis (c, ZMB 76699; 1750–2200 m asl).
Tadpoles were collected with nets and preserved in 8% formalin. Pieces of the tail were stored in 96% ethanol. Adults of all three species have been collected on Mount Manengouba and liver samples of these were taken for genetic comparisons. All voucher specimens were deposited at the Museum für Naturkunde Berlin (ZMB; Appendix 1). Tadpoles were identified using DNA-barcoding, analyzing approximately 490 bp of the mitochondrial 16S rRNA (see Rödel et al. 2009 for methods).
Accepted by M. Vences: 2 Dec. 2011; published: 25 Jan. 2012
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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. The descriptions of tadpoles are based on the genotyped specimens and further individuals of the same site and comparable developmental stage. Measures in the text are presented as: first the genotyped specimens followed by measures of the other tadpoles (all measures in mm). Developmental stage was determined following Gosner (1960). The morphological terminology is based on Altig & McDiarmid (1999). Measurements were taken by one person (MH) with dial callipers or with the aid of a dissecting microscope (accuracy ± 0.1 mm). Photographs of tadpoles are multifocus-photos, taken with a Leica® DFC420 digital camera on a Leica® MZ 95 dissecting microscope. Single pictures were combined using the Automontage® software version 5.03.0061 (Syncroscopy). All tadpoles could be assigned to one of two morphological groups. Each morphological group originated from one particular river site only. By comparing the sequences of the tadpoles to the adult toads, tadpoles of the two morphological groups could be determined as W. tandyi (intraspecific sequence divergence 0.00-0.41%; N= 4 incl. DPL5107, see below), and W. bambutensis (intraspecific sequence divergence 0.00%; N= 3), respectively. The interspecific sequence divergence of the three species ranged from 5.93-7.74%, thus confirming specific tadpole affiliation. A W. „mertensi“ sequence available at GenBank (DQ283348; specimen: DPL5107) was almost identical to our W. tandyi sequences (Appendix 1) and thus likely is based on a misidentified toad. Werneria bambutensis (Fig. 2b, d, f).– Three tadpoles were collected at 2135 m asl. The medium sized river (N5°00.589', E9°51.414') was flowing through a fragment of secondary forest, the vegetation was degraded by cattle. A waterfall was close by. The W. bambutensis tadpoles were encountered attached to stones in the middle of the river, where the water current was the strongest. In parts of the river course, where the water current was slower we caught Phrynobatrachus tadpoles and the tadpoles of Cardioglossa manengouba Blackburn, 2008. All three tadpoles (ZMB 76706/76705/76707; total length: -/18.5/13.2) were in Gosner stage 25. The body is robust and broad, less flattened than in W. tandyi; in dorsal view the body shape is egg-like, the apical part of the egg being the posterior end of the body; snout broadly rounded but more pointed in dorsal view than in W. tandyi (Fig. 2b, d); body longer (5.4/6.9/5.0) than broad (3.7/5.1/3.5), body width reaching 69-74% of body length, maximum body width on level with eyes; body length reaches 36-37% of total length; body higher than in W. tandyi, height (2.2/3.6/2.2) ranges between 41-52% of body length; nostrils small and comma shaped, positioned dorsally, in two tadpoles, they are closer to eyes (distance: 0.7/1.2/0.8) than to snout tip (distance: 1.0/1.5/0.7), in one tadpole nostrils are closer to the snout; snout-nostril distance ranges between 14-22% of body length; eyes are positioned dorsolaterally, closer to snout (distance: 1.8/2.6/1.6) than to vent, eye diameter (0.4/0.6/0.5) 7-10% of body length; interorbital distance (1.5/2.1/1.6) exceeds the internarial distance (1.0/1.3/0.9) by the factor 1.5-1.8; relatively short tail (-/12.2/8.1) with narrow fin; tail length 1.6-1.8 of body length; dorsal fin originating at the base of the tail; ventral fin originating just marginally posterior to the vent opening; fin height along first two thirds higher in dorsal part, higher in ventral part at last third of tail; fin tip broadly rounded; maximum tail height including fins (2.9/3.6/2.6) equals or passes body by 1.31.4; the tail axis broad (1.2/1.8/1.1) and muscular, narrowing towards tail tip after about half of tail length; maximum height of tail axis (2.0/2.3/1.8) between 64-69% of total tail height; median vent tube; spiraculum sinistral (distance: posterior corner of eye to posterior end of spiraculum: 1.8/2.7/1.9), barely visible in dorsal view, originating at the last third of body; spiraculum tube very short (0.4/0.9/0.3) and narrow, 6-12% of body length; mouth opens ventrally; oral disc broad, sucker-like (Fig. 2f), oral disc width equals body width; labial tooth row formula is 2/3; upper jaw narrow, heavily keratinized part forms a compressed ‘m’; lower jaw a very flat, narrow V; anterior lip widely rounded with a median gap, without papillae, margin smooth; anterior lip carrying supra-angular teeth rows; small uniserial marginal papillae; posterior lip consists of two wide, rounded dermal lobes, covered densely with small papillae; dorsal colour almost uniform brownish; only upper lip grey; a clear transverse band behind eyes much less pronounced than in W. tandyi; spiraculum brownish, with the opening translucent in appearance; venter speckled with brown dots; tail axis light brown, clearer than body; fin greyish without pattern. Werneria tandyi (Fig. 2a, c, e).—Three tadpoles were collected in a medium sized river at 1319 m asl (N5°05.439', E9°48.406'). The collection site was within a fragment of secondary forest, which is frequently invaded by cattle. The forest fragment is surrounded by areas dominated by small-scale subsistence farming. The W. tandyi tadpoles were collected attached to stones in fast flowing parts of the river. Other tadpoles inhabiting this micro-habitat were those of Trichobatrachus robustus. The genotyped individual (ZMB 76694) is in Gosner stage 30. Two further individuals are in stage 28 (ZMB 76695/ 76696). The body (total length: -/22.5/23.6) is dorsoventrally flattened; robust and broad, almost rectangular shaped in dorsal view (Fig. 2a, c); body longer (9.4/8.0/9.0) than broad (6.9/6.0/6.5), body width reaching 72-75% of body length, maximum width reaches just posterior to eyes; body length reaches 36-38% of total length; body height (4.5/3.2/4.0) ranges between 4048% of body length; snout broadly rounded in dorsal view; nostrils small and round, positioned dorsally, closer to eyes (distance: 1.6/1.4/1.4) than to snout tip; snout-nostril distance (2.3/1.9/2.3) ranges between 24-26% of body length; eyes are positioned dorsally, almost at mid-body (distance to snout: 4.0/3.4/3.8); eyes small (diameter: 0.8/0.7/0.8), eye diameter 9% of body length; interorbital distance (2.6/2.3/2.4) exceeds the internarial distance (1.7/1.5/1.7) by 1.4-1.5; relatively short tail (-/ 14.5/14.5) with narrow fin; tail length 1.6-1.8 of body length; dorsal fin originating after about 1/3 of tail length; ventral fin originating at about half of tail length; fin height almost equal in dorsal and ventral fin; fin tip broadly rounded; maximum tail height including fins (3.8/3.5/3.6) ranges between 84-109% of body height; the tail axis broad (2.8/2.0/2.1) and muscular, narrowing towards tail tip after about half of tail length; maximum height of tail axis (2.7/2.3/2.6) between 66-72% of total tail height; median vent tube; spiraculum sinistral (distance: posterior corner of eye to posterior end of spiraculum: 3.5/3.3/3.0), visible in dorsal view, originating at the last third of body just behind broadest part of body; spiraculum tube short (0.9/0.7/0.5), 6-10% of body length; mouth opens ventrally; oral disc broad, sucker-like (Fig. 2e), oral disc width almost reaches body width;
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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.
FIGURE 2. Lateral, dorsal and mouthpart views of Werneria tadpoles. Shown are a Gosner stage 28 tadpole of W. tandyi (a, c, e; ZMB 76696) and a Gosner stage 25 tadpole of W. bambutensis (b, d, f; ZMB 76705); scale bars = 2 mm.
labial tooth row formula is 2/3; upper jaw narrow, heavily keratinized part forms a compressed ‘m’; lower jaw medially deeply notched, keratinized parts oval shaped and separated; anterior lip widely rounded, expanded without papillae, margin smooth;
WERNERIA TADPOLES FROM CAMEROON
Zootaxa 3172 © 2012 Magnolia Press ·
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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. anterior lip carrying supra-angular teeth rows; small uniserial marginal papillae; posterior lip consists of two wide, rounded dermal lobes, the posterior one with a narrowly serrated margin, margin of anterior one with a smooth edge; dorsal colour brownish; anterior part of body greyish light, almost translucent; a pair of clear blotches between nares and eyes; a clear transverse band behind eyes; spiraculum translucent; venter translucent; tail axis light brown, clearer than body; fin greyish and translucent without pattern. Werneria tadpoles, including the two species described herein, are exotrophic, lotic and suctorial, thus being similar to other torrenticolous anuran tadpoles, e.g. the neotropical Atelopus or south-east Asian Ansonia (Altig & McDiarmid 1999). The W. bambutensis tadpoles confirm Amiet’s (1976) statement that tadpoles of this species are more robust than other congeners. His notes of dark brown tadpoles with a tail speckled with golden chromatophores (Amiet 1972), indicate that the colour might vary. However, body form, insertion site of tail fin and characters of the oral disc, such as shape and keratinisation of jaw sheaths, allow for correct species identification. Differences of the tadpoles described as W. tandyi herein, to the description of W. tandyi tadpoles by Rödel et al. (2004), indicate an incorrect assignment of the latter tadpoles to this species. Their body shape was similar to the W. tandyi tadpoles identified by barcoding, they likewise had a body which was much flattened dorsoventrally. In contrast, their jaw sheaths are more similar to those of W. bambutensis, i.e. the keratinized lower jaw is not consisting of two parts. It therefore seems likely that these tadpoles, collected by J.-L. Perret at Nsoung, Mount Manengouba and inventoried in Geneva as W. tandyi (MHNG 1020.44-1 & 2) are W. mertensiana given that this is the only other Werneria species that occurs at this location. However, this remains to be confirmed. The tadpoles of W. preussi, described by Mertens (1938) have oral discs similar to W. bambutensis but differ from those by a much more rounded and flatter body.
Acknowledgements We thank the Cameroonian Ministry of Forestry and Wildlife (MINFOF) for issuing research, collection and export permits. Nono L. Gonwouo is thanked for the invaluable help given during the planning of the field work. We thank our local field assistants for their help during data collection. MH was supported by scholarships from the Federal State of Berlin (Elsa-Neumann-Stipendium) and the German Academic Exchange Service (DAAD). Eureka! provided the camping equipment. Further financial support was provided by the Stadtgruppe Leipzig of the German Society of Herpetology (DGHT), and the newspaper “Seesener Beobachter”. SPL is supported by a Swiss National Science Foundation grant (31003A-133067).
Literature cited Altig, R. & McDiarmid, R.W. (1999) Tadpoles: the biology of anuran larvae. The University of Chicago Press, Chicago, London. Amiet, J.-L. (1972) Description de trois Bufonidés orophiles du Cameroun appartenant au groupe de Bufo preussi Matschie (Amphibiens Anoures). Annals de la Faculté des Sciences du Cameroun, 11, 121–140. Amiet, J.-L. (1976) Observations anatomiques et biologiques sur le genre Werneria Poche, 1903. Revue de Zoologie Africaine, 90, 33–45. Gosner, K.L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16, 183–190. Mertens, R. (1938) Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der senckenbergischen naturforschenden Gesellschaft, 442, 1–52. Rödel, M.-O., Schmitz, A., Pauwels, O.S.G. & Böhme, W. (2004) Revision of the genus Werneria Poche, 1903, including the description of two new species from Cameroon and Gabon (Amphibia: Anura: Bufonidae). Zootaxa, 720, 1–28. Rödel, M.-O., Sandberger, L., Doumbia, J. & Hillers, A. (2009) Revalidation of Phrynobatrachus maculiventris Guibé & Lamotte, 1958 and description of its aposematic coloured tadpole. African Journal of Herpetology, 58, 15–27.
Appendix 1. Werneria vouchers from Mount Manengouba, Cameroon and GenBank accession numbers. Werneria bambutensis: ZMB 76705-76707 (tadpoles; GenBank # of ZMB 76706: JN569200), N5°00.589', E9°51.414', 2135 m asl, 7 November 2010, medium sized river, secondary forest; ZMB 76698, N05°00.963', E09°52.124', 2096 m asl, 16 September 2010, leaf litter in forest; ZMB 76699-76704, N05°01.074’, E09°51.945’, 2130 m asl, 5 August 2008, leaf litter in forest; ZMB 76851-76855 (GenBank # of ZMB 76853: JN569198; of ZMB 76854: JN569199), N05°01’, E009°52’, 2096 m asl, 8 October 2007, secondary forest. Werneria mertensiana: ZMB 76693, N04°58.773’, E09°53.820’, 1259 m asl, 9 August 2008, small fast flowing river, leaf litter in secondary forest; ZMB 76856 (GenBank # JN569197), N04°57', E009°51', 1106 m asl, 16 November 2006. Werneria tandyi: ZMB 76694-76696 (tadpoles, GenBank # of ZMB 76694: JN569194), N5°05.439', E9°48.406', 1319 m asl, 5 December 2010, medium sized river, secondary forest; ZMB 76697 (GenBank # JN569195), N04°58.773’, E09°53.820’, 1259 m asl, 9 August 2008, small fast flowing river, leaf litter in secondary forest; ZMB 76857, N04°59', E009°52', 1525 m asl, 10 March 2006; ZMB 76858, N04°58', E009°52', 1375 m asl, 10 April 2006; ZMB 76859-76860 (GenBank # of ZMB 76860: JN569196), N05°01’, E009°46’, 1480 m asl, 17 December 2006.
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