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Title Taxonomic Relationships of Ansonia Anotis Inger, Tan

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Title Taxonomic Relationships of Ansonia Anotis Inger, Tan View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Kyoto University Research Information Repository Taxonomic Relationships of Ansonia anotis Inger, Tan, and Title Yambun, 2001 and Pedostibes maculatus (Mocquard, 1890), with a Description of a New Genus (Amphibia, Bufonidae) Author(s) Matsui, Masafumi; Yambun, Paul; Sudin, Ahmad Citation Zoological Science (2007), 24(11): 1159-1166 Issue Date 2007-11 URL http://hdl.handle.net/2433/85317 Right (c) 日本動物学会 / Zoological Society of Japan Type Journal Article Textversion publisher Kyoto University ZOOLOGICAL SCIENCE 24: 1159–1166 (2007) © 2007 Zoological Society of Japan Taxonomic Relationships of Ansonia anotis Inger, Tan, and Yambun, 2001 and Pedostibes maculatus (Mocquard, 1890), with a Description of a New Genus (Amphibia, Bufonidae) Masafumi Matsui1*, Paul Yambun2 and Ahmad Sudin3 1Graduate School of Human and Environmental Studies, Kyoto University, Sakyo, Kyoto 606-8501, Japan 2Research and Education Division, Sabah Parks, P.O. Box 10626, Kota Kinabalu 88806, Sabah, Malaysia 3Institute for Tropical Biology and Conservation, University Malaysia Sabah, Teluk Sepanggar, Locked Bag 2073, Kota Kinabalu 88999, Sabah, Malaysia Examination of types and recently collected specimens revealed that Ansonia anotis Inger, Tan, and Yambun, 2001 and Pedostibes maculatus (Mocquard, 1890), both described from Kinabalu, Sabah, Malaysia, are hardly differentiated morphologically. Analyses of a total of 2,427 bp of the 12S rRNA, tRNAval, and 16S mitochondrial rRNA genes revealed that the two species are very close genetically. Thus A. anotis is regarded as conspecific and is synonymized with P. m a c u l a t u s . Genetically, this species proved to form a lineage distinct from other bufonids from Southeast Asia, including spe- cies of Ansonia and Pedostibes. Because the species has also some unique morphological traits different from known bufonid genera, we propose to establish a new genus for Nectophryne maculata Mocquard, 1890. Key words: Malaysia, molecular phylogeny, new genus, Sabah, synonymy but Pelophryne is small in body size, and lays small num- INTRODUCTION bers of large yolky eggs, while Pedostibes is moderate to Southeast Asia is one of the centers of amphibian diver- large sized, and its eggs are numerous and small. Recent sification (Inger, 1999), and this holds for the toad family molecular studies indicate close relationships of Pedostibes Bufonidae. On the island of Borneo, as many as six bufonid with Bufo asper Gravenhorst, 1829 and B. juxtasper Inger, genera (Bufo Laurenti, 1768, Ansonia Stoliczka, 1870, 1960 (Frost et al., 2006; our own observations, see Results). Leptophryne Fitzinger, 1843, Pedostibes Günther, 1876, Inger et al. (2001) described Ansonia anotis Inger, Tan, Pelophryne Barbour, 1938, and Pseudobufo Tschudi, 1838) and Yambun, 2001 from Sayap, in the Kinabalu National have been recorded (Inger, 1966; Inger and Tan, 1996; Park of Sabah, Malaysian part of northern Borneo (Fig. 1). Malkmus et al., 2002), and the number is even greater if we The species is characterized mainly by lack of a tympanum, admit Frost et al.’s (2006) proposal to split Bufo into several large spatulate finger disks, and the unique morphology of distinct genera. This might promote uncovering the para- the putative larva. However, because only several speci- phyletic nature of the genus Bufo (e.g., Graybeal and mens have been obtained since its description, details of the Cannatella, 1995), but applying unfamiliar names for many species are unclear. Recently, on the joint expedition of taxa, without substantially providing their diagnostic charac- UMS (University Malaysia Sabah) and JICA (Japan Interna- ters, could result in further taxonomic confusion. tional Cooperation Agency) to the Crocker Range National Some other Bornean genera are taxonomically more Park (Fig. 1), Sabah, in 2002, four toads were collected. conservative than Bufo and occupy specific ecological These specimens were very similar to Pedostibes maculatus niches (Inger, 1958). Of these, Ansonia is characterized by (Mocquard, 1890), originally described as Nectophryne, and small to medium adult body size and a unique larva with a were identified as that species (Kueh et al., 2004). Like A. large oral sucker adapted for life in torrents. Both anotis, P. maculatus is characterized by lack of the tympa- Pelophryne and Pedostibes are adapted for arboreal life, num and large finger disks (Boulenger, 1918; Inger, 1966), and these species could not be clearly distinguished from one another by any morphological traits described in the lit- * Corresponding author. Phone: +81-75-753-6846; Fax : +81-75-753-6846; erature. E-mail : [email protected] We compared the two species morphologically using doi:10.2108/zsj.24.1159 available specimens, including three syntypes of N. maculata, 1160 M. Matsui et al. Fig. 1. Map of Borneo showing the localities where Ansonia anotis (open circle) and Pedostibes maculatus (open square) were recorded. 1, Sayap; 2, Kiau; 3, Sungei Purulon; 4, Trail 5 of Ulu Kimanis. Hatched areas show the Kinabalu and the Crocker Range National Parks (KNP and CRNP, respectively). stored in the collection of Museum National d’Histoire length (FL). All measurements were made to the nearest 0.1 mm Naturelle, Paris (MNHNP), to determine their taxonomic with dial calipers. Dimensions were converted to a percentage ratio relationships. Further, in order to clarify their phylogenetic (R) in relation to SVL for comparisons, although small sample sizes relationships among Southeast Asian bufonids, we analyzed examined prohibited statistical comparisons. We made slight dis- sequences of the 12S and 16S mitochondrial rRNA genes sections to examine tympanic structure and gonads, and prepared radiographs to examine gross osteology. of the two species, as well as of representatives of related For genetic comparisons, we studied one specimen each of A. bufonids for comparisons. anotis and P. maculatus from among the specimens used for mor- MATERIALS AND METHODS phological comparisons. For comparisons, we used the following species (Table 1): A. malayana Inger, 1960; A. hanitschi Inger, For morphological comparisons, we studied three specimens of 1960; A. longidigita Inger, 1960; A. fuliginea (Mocquard, 1890); Ansonia anotis (Sabah Parks, Kota Kinabalu [SP] 01762 (holotype) Pedostibes hosii (Boulenger, 1892); P. rugosus Inger, 1958; from Sungei (River) Wario, Sayap, Mt. Kinabalu, Sabah; SP 26033 Pelophryne misera (Mocquard, 1890); P. brevipes (Peters, 1867); and 26043 from Riou Hutan Simpan, Kiau, Mt. Kinabalu, Sabah), Leptophryne borbonica (Tschudi, 1838); Bufo juxtasper (=Phrynoidis three syntypes of N. maculata from Kinabalu (MNHNP 1899-266– juxtaspera in Frost et al., 2006); B. melanostictus Schneider, 1799 268; syntypes), and two of four specimens of P. maculatus (Grad- (=Duttaphrynus melanostictus in Frost et al., 2006); B. divergens uate School of Human and Environmental Studies, Kyoto University Peters, 1871 (=Ingerophrynus divergens in Frost et al., 2006); [KUHE] 38505 and 38506 from Ulu Kimanis of the Crocker National Didynamipus sjostedti Andersson, 1903 from GenBank Park [Trail 5 of UMS 2002 expedition to the Crocker Range], (AY325991); and Atelopus flavescens Duméril and Bibron, 1841 Sabah). from GenBank (DQ283259). We chose Dendrobates auratus Six body measurements were taken, following Matsui (1984): (Girard, 1855) from GenBank (AY326030), a member of the sup- 1) snout-vent length (SVL), 2) head width (HW), 3) lower arm length posed bufonid sister family, Dendrobatidae Cope, 1865 (Frost et al., (LAL), 4) thigh length (THIGH), 5) tibia length (TL), and 6) foot 2006), as outgroup (Table 1). Taxonomy of a Bornean Toad 1161 Table 1. Samples used for DNA analyses and GenBank accession numbers. *BORNEENSIS=Institute for Tropical Biology and Conservation, University Malaysia Sabah; KUHE=Graduate School of Human and Environmental Studies, Kyoto University; SP=Sabah Parks. Species Voucher* Locality Accession number Ansonia anotis SP 26033 Kiau, Mt. Kinabalu, Sabah AB331708 Ansonia fuliginea KUHE 17537 Mt. Kinabalu, Sabah AB331709 Ansonia hanitschi BORNEENSIS 22640 Silau Silau, Mt. Kinabalu, Sabah AB331710 Ansonia longidigita BORNEENSIS 12463 Mahua, Crocker Range, Sabah AB331711 Ansonia malayana KUHE 15472 Larut, Peninsular Malaysia AB331712 Bufo juxtasper KUHE 12363 Bareo, Sarawak AB331713 Bufo melanostictus KUHE 10524 Marudi, Sarawak AB331714 Bufo divergens BORNEENSIS 09169 Tawau Hills NP, Sabah AB331715 Leptophryne borbonica BORNEENSIS 08127 Ulu Kimanis, Crocker Range, Sabah AB331716 Pedostibes hosii BORNEENSIS 22088 Tawau Hills NP, Sabah AB331717 Pedostibes maculatus BORNEENSIS 08425 Ulu Kimanis, Crocker Range, Sabah AB331718 Pedostibes rugosus SP 21556 Mahua, Crocker Range, Sabah AB331719 Pelophryne brevipes KUHE 35585 Peninsular Malaysia AB331720 Pelophryne misera KUHE 37191 Mt. Kinabalu, Sabah AB331721 Didynamipus sjostedti – Cameroon AY325991 Atelopus flavescens – French Guiana DQ283259 Dendrobates auratus – – AY326030 We extracted DNA from small amounts of frozen or ethanol- with 1,000 replicates (Hedges, 1992). We regarded tree topologies preserved tissues using standard phenol-chloroform extraction pro- with bootstrap values (BS)≥70% as sufficiently resolved cedures (Hillis et al., 1996). We conducted amplifications by the (Huelsenbeck and Hillis, 1993). polymerase chain reaction (PCR) with the primers Thr Lrn (AAAR- We also performed Bayesian analyses using the
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