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Larval Identities of Ansonia Hanitschi Inger, 1960 (Amphibia: Bufonidae

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Larval Identities of Ansonia Hanitschi Inger, 1960 (Amphibia: Bufonidae Larval identities of Ansonia hanitschi and Polypedates colletti SALAMANDRA 44 2 85-100 Rheinbach, 20 May 2008 ISSN 0036-3375 Larval identities of Ansonia hanitschi Inger, 1960 (Amphibia: Bufonidae) and Polypedates colletti (Boulenger, 1890) (Amphibia: Rhacophoridae) from East Malaysia (Borneo) Alexander Haas & Indraneil Das Abstract. We describe the tadpoles of Ansonia hanitschi Inger, 1960 and Polypedates colletti (Boul- enger, 1890) from East Malaysia, Borneo. The morphological description is supplemented by photos of living specimens and SEM images of external and internal features. Identification was based on morpho- logical and genetic characters (16S rRNA). The tadpole of A. hanitschi is among the largest tadpoles de- scribed for the genus Ansonia from Borneo. This suctorial-rheophilous tadpole has been collected from fast-flowing rocky streams, in association with foaming currents. The tadpole of P. colletti is redescribed due to uncertainties about its identity and lack of photographic evidence in previous descriptions. It is an inhabitant of stagnant waters, living in and on flooded leaf litter accumulations. Key words. Bufonidae, Rhacophoridae, Ansonia hanitschi, Polypedates colletti, tadpole description, larva, larval morphology, oral disk. Introduction cially Inger (1966, 1983, 1985, 1992) and In- ger et al. (2006). In a recent study (Das & At present 5,574 anuran species have been de- Haas 2006), we summarized the literature scribed (www.amphibiaweb.org). New spe- that provides data on larval identities of Bor- cies continue to be discovered from even nean amphibians. According to this survey, well-explored regions of the earth, but the in- approximately 55% of the Bornean amphib- ventory and description of larval forms of the ian fauna have known larvae; since then, known species are far from complete. How- one new description has become available ever, recognition of the larval stages and de- (Inger et al. 2006). Even when descriptions scriptions of tadpoles are essential for many are available, these are often superficial or in purposes and research questions, such as sur- abbreviated form, sometimes without draw- veys, habitat inventories, studies on resource ings, often without images (photographs), use in habitats, interspecific competition preventing unequivocal identification of lar- studies, and conservation efforts. Tadpoles vae in the field or laboratory and the repro- are so different anatomically and ecologically duction of the former. Furthermore, the de- from the adult stage that they deserve sepa- scriptions surveyed by Das & Haas (2006) rate treatment (McDiarmid & Altig 1999). were frequently without deposited well-doc- Interest in tadpoles has increased in the re- umented voucher specimens, making identi- cent past, and various studies have attempt- ties of described tadpoles uncertain. This is ed to summarize regional tadpole faunas (for especially a problem with old species names example, Chou & Lin 1997, Leong & Chou that are now considered to contain two or 1999, Anstis 2002). more valid species, e.g., Leptobrachium mon- For the island of Borneo, and especially the tanum Fischer, 1885 and Hylarana signata East Malaysian states of Sabah and Sarawak, (Günther, 1872) (see Malkmus et al. 2002, Robert F. Inger laid a solid foundation for Brown & Guttman 2002). tadpole research in a series of papers, espe- The present work describes two larval © 2008 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT) http://www.salamandra-journal.com 85 Alexander Haas & Indraneil Das forms: the tadpoles of Ansonia hanitschi In- from his earlier account (Inger 1966). This ger, 1960 (Bufonidae) and Polypedates col- uncertainty and the lack of drawings or pho- letti (Boulenger, 1890) (Rhacophoridae). tographs in Inger’s accounts warrant a re-de- Ansonia hanitschi is a stream toad endemic scription of the P. colletti tadpole. to Borneo. Adults have been reported from streams within submontane and montane forests above 950 m above sea level (Inger & Materials and methods Stuebing 2005). The species is known from Gunung Kinabalu and the adjacent Crock- Tadpoles of A. hanitschi were collected at er Range (Sabah), Gunung Mulu (northern Sungei Silau-Silau, Gunung Kinabalu Na- Sarawak) and from the north-eastern parts tional Park (N 06°00.639’, E116°32.418’; 1,450 of Kalimantan (Malkmus et al. 2002, Inger m asl), Sabah. Calling adults and breeding & Stuebing 2005). The tadpoles of A. han- congregations of A. hanitschi were present itschi were mentioned briefly inMalkmus et in large numbers at the collection sites at the al. (2002) and Inger & Stuebing (2005), de- time of collecting. No adults of other Ansonia scribing these tadpoles as ”... black and tear- species were encountered at Sungei Silau-Si- drop shaped. Maximum length is 20 mm.” lau during the two seasons of collecting. One This description is so general that it could fit adult voucher was collected from Mesilau several Ansonia species and it conflicts with (also within the Gunung Kinabalu massif, ca. our findings. Hence we doubt that this de- 2,000 m asl; voucher ZRC 1.11911; ex ID 8161) scription is valid for A. hanitschi and con- and tissue was taken for DNA sequencing. sider the tadpole of A. hanitschi as formally Tadpoles of P. colletti were collected in undescribed. Sarawak at (1) Sama Jaya Nature Reserve (N Polypedates colletti (Boulenger, 1890) is 01°31.258’, E110°23.248’) from a leaf litter filled a widespread species in lowland dipterocarp shallow depression next to the trail (field forests, alluvial forests, and peat swamps. It number: 189), (2) Gunung Santubong (field shares its habitat preference with, for example, number: 135), Summit Trail, from leaf litter Ingerophrynus quadriporcatus (Boulenger, filled depression next to trail, and (3) Gu- 1887), Hylarana baramica (Boettger, 1900) nung Mulu National Park, Camp 5, Keran- and H. glandulosa (Boulenger, 1882), and is gas Forest Trail, approx. 50 m off trail, pool often found in syntopy with these species. The from an uprooted tree in Kerangas forest veg- inaccessibility of the swamps in lowland Bor- etation (field number: 384). Living tadpoles neo and the difficulties in finding tadpoles in were photographed and subsequently anaes- the dark brown, leaf litter filled waters may thetized and preserved in either 4% neutral- be reasons why the tadpoles of swamp spe- buffered formalin or absolute ethanol.A dults cies have rarely been collected and described; from two sites were sampled for liver and only recently were the tadpoles of I. quadri- muscle tissue (absolute alcohol): one adult porcatus (Leong & Chou 1999) and H. glan- from Loagan Bunut National Park (ZRC dulosa (Inger et al. 2006) described from 1.11912, ex-ID 8094) and one from Mulu Na- such habitats. Inger (1966) described the tional Park, Camp 5, Deer Pond (ZRC 1.11914, larva of P. colletti, however, without provid- ex ID 8571). ing drawings or photos. Later, Inger (1985: Both species in this study were identi- 71) concluded for other samples that some fied from a series of metamorphic and post- ”.. factors leave the identification uncertain”. metamorphic specimens and DNA sequence Also Inger (1985) described samples from matching. Metamorphs of A. hanitschi had Sabah and Sarawak as differing markedly in slightly dilated finger tips. In metamorphs, tail tip shape and colouration, and deviating the tip of the first finger did not reach the 86 Larval identities of Ansonia hanitschi and Polypedates colletti Fig. 1. Colouration of larval Ansonia hanitschi in life (various specimens): a) lateral view of fully grown tadpole (stage 38), note the short acuminate tip; b) dorsal view of metamorph (stage 41) with emerging juvenile dorsal pattern on dorsum and legs; c) adult individual from Sg. Silau-Silau; d) metamorph; e) ventral view, note the length of upper lip keratodont rows curving laterally around lower lip keratodont rows, LTRF 2/3, widely separate upper jaw sheaths; f hand (ventral view) of stage 42 specimen. 87 Alexander Haas & Indraneil Das disk of the second when adpressed (Inger, bp, 41 non-matching sites) to 100% (EF624065 1966) and the typical adult dorsal pattern was vs. EF624066). A Genbank BLAST search present with congruence in even subtle de- yielded P. colletti (AF215354.1, by M. Vences) tails of the patterns on the head and shoulder as best match, differing from EF566974 in 7 parts (Fig.1). base pairs in the overlapping 528 bp region Genetic matching of tadpoles with adults (98.7%). Additional tadpole voucher speci- was based on the partial mitochondrial 16S mens were deposited at the Zoological Mu- rRNA gene. DNA was extracted from mac- seum, Hamburg (ZMH, see Table 1). erated muscle or liver tissue according to We apply the classification ofFrost (2007) standard methods (Hillis et al. 1996) and based on suggestions of Frost et al. (2006). was stored at -20°C. DNA amplification of We used terms for tadpole descriptions from partial 16S rRNA gene sequences was done Inger (1985), Altig & McDiarmid (1999), with peqGOLD PCR-Master-Mix Y (Peqlab) and Anstis (2002). Terminology for inter- according to the manufacturer’s guide- nal oral features was adopted from Wasser- lines. The sense primer (16SC) 5’- GTRG- sug (1976). Tadpole stages were recorded in GCCTAAAAGCAGCCAC - 3’ and the anti- using Gosner’s table (1960). Measurements sense primer (16SD) 5’- CTCCGGTCT- were taken from digital images using Im- GAACTCAGATCACGTAG - 3’ were chosen ageJ software (National Institute of Health). (Rafe Brown pers. comm).
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