Zootaxa, A new species of Gastrophrynoides - La Sierra University

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use.Commercial sale or deposition in a public library or website is prohibited.Zootaxa 2124: 63–68 (2009)www.mapress.com/zootaxa/Copyright © 2009 · Magnolia PressArticleISSN 1175-5326 (print edition)ZOOTAXAISSN 1175-5334 (online edition)A new species of Gastrophrynoides (Anura: Microhylidae): an addition to apreviously monotypic genus and a new genus for Peninsular MalaysiaCHAN KIN ONN 1,5 , L. LEE GRISMER 1, 3 , NORHAYATI AHMAD 1, 2 & DAICUS BELABUT 1, 41Institute for Environment and Development (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor Darul Ehsan,Malaysia2School of Environment and Natural Resource Sciences, Faculty of Science and Technology, Universiti Kebangsaan Malaysia, 43600Bangi, Selangor Darul Ehsan, Malaysia. E-mail: yati_68@yahoo.co.uk3Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California, 92515-8247 USA.E-mail: lgrismer@lasierra.edu4Institute of Biological Sciences,Faculty of Science, University of Malaya, 50603 Kuala Lumpur, Malaysia.E-mail: daicus@um.edu.my5Corresponding author. E-mail: kin_onn@yahoo.comAbstractA new species of microhylid frog of the genus Gastrophrynoides is described from Gunung Besar Hantu, in the state ofNegeri Sembilan, adding a new generic record to the family Microhylidae of Peninsular Malaysia and an additionalspecies to the previously monotypic genus Gastrophrynoides. This new species can be distinguished from its onlycongener, G. borneensis by having an immaculate, grayish brown dorsum (instead of a spotted one); a longer snout(2.6–3.0 vs. 2.0–2.5 times diameter of eye); and a single, large, oval outer metacarpal tubercle beneath the hand (insteadof smaller, paired, rectangular outer metacarpal tubercles in G. borneensis).Key words: Gunung Besar Hantu, Gastrophrynoides borneensis, Negeri Sembilan, frogIntroductionGastrophrynoides Noble 1926, as currently constituted, is a monotypic genus containing the species G.borneensis (Boulenger 1897). This rare and elusive frog is endemic to Borneo, occurring in Sarawak (Inger1966) and Sabah (Kueh & Sudin 2008). A single specimen recorded from Sumatra, Indonesia is doubtful andrequires further confirmation before it is accepted (Inger 1966).Frost et al. (2006), without comment, removed Gastrophrynoides from the subfamily Microhylinae butaddressed it no further. Subsequent molecular studies have not addressed this taxon (Van der Meijden et al.2007) due to its rarity, leaving its phylogenetic placement unresolved. Knowledge of the ecology of this frogremains equally limited due to the scarcity of findings. Previous accounts of G. borneensis were all fromlowland forests less than 500 m a.s.l (Inger & Stuebing 2005), where frogs were found beneath dead leaves onthe forest floor or in water-filled holes. More recently, Kueh & Sudin (2008) found a specimen inside a rottinglog in a drying stream bed at 878 m elevation in Tawau Hills Park, Sabah.In April 2008, we obtained three specimens of a strange frog from the top of Gunung Besar Hantu, NegeriSembilan, Peninsular Malaysia at 800–1400 m elevation (Figure 1). We assigned them to the familyMicrohylidae owing to their lack of maxillary teeth; lack of parotoid glands; and a firmisternal pectoral girdlewith non-overlapping epicoracoids; well developed coracoids reaching the midline of the girdle and scapulae;a large, cartilaginous sternum; and no clavicles or osmosternum. We identified them as belonging to the genusGastrophrynoides on the basis of having completely smooth skin and a long, pointed snout more than twiceAccepted by M. Vences: 15 May 2009; published: 3 Jun 2009 63

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use.Commercial sale or deposition in a public library or website is prohibited.the diameter of the eye. However, these frogs differ from G. borneensis in having an immaculate, grayishbrown dorsum as opposed to a spotted one, a longer snout (2.6–3.0 vs. 2.0–2.5 times diameter of eye), andhaving a single, large, oval outer metacarpal tubercle beneath the hand as opposed to smaller, paired,rectangular outer metacarpal tubercles in G. borneensis. Owing to a combination of differences inmorphology, color-pattern and geographic distribution, we evaluate this population within the framework of aGeneral Lineage Species Concept (De Queiroz 1998) and use morphological diagnosibility and geographicisolation (e.g. Grismer 1999; McGuire et al. 2007) as criteria for lineage independence. Given that this newspecies of microhylid was found on an isolated mountain top on the west coast of Peninsular Malaysia,whereas G. borneensis, from which it is morphologically different, is restricted Borneo, there is no doubt thatthese two allopatric forms represent independent evolutionary lineages, and as such, we describe thePeninsular Malaysian population as a new species.FIGURE 1. Type locality of Gastrophrynoides immaculatus and distribution of G. borneensis.Materials and methodsSpecimens were photographed and liver tissue was taken prior to preservation and stored in 95% ethanol.Specimens were fixed with 10% formalin and stored in 70% ethanol. Measurements were taken withMitutoyo digimatic caliper to the nearest 0.1 mm. Radiographs were prepared using a Faxitron closed x-raysystem with an exposure time of 20 sec at 20 kV.Images were recorded on Polaroid type 55 film.Measurements taken: snout-vent-length (SVL), taken from tip of snout to vent; head length (HL), measuredfrom the posterior margin of lower jaw to tip of snout; head width (HW), taken immediately posterior to eyes;eye diameter (ED), distance between anterior and posterior corners of upper and lower eyelids; width of uppereyelid (ELW), measured from the base of upper eyelid to lateral edge at its widest point; interorbital distance(IOD), distance across top of head between medial margins of orbits at their closest points; snout length (SL),measured from anterior corner of the orbit to tip of snout; distance of eye to nostril (DETN), measured fromanterior corner of eye to posterior margin of nostril; distance of nostril to tip of snout (DNTS), measured fromanterior margin of nostril to tip of snout; internarial distance (IND), measured from medial inner margins ofnostrils; tympanum diameter (TD), horizontal width of tympanum at its widest point; tibia length (TiL),measured from center of knee to center of ankle. All specimens have been deposited at the UniversitiKebangsaan Malaysia Herpetological Collection (HC), Bangi, Malaysia and La Sierra UniversityHerpetological Collection (LSUHC), La Sierra University, Riverside, California, USA. FMNH refers to theField Museum of Natural History, Chicago, USA.64 · Zootaxa 2124 © 2009 Magnolia PressCHAN ET AL.

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use.Commercial sale or deposition in a public library or website is prohibited.SystematicsGastrophrynoides immaculatus sp nov.Figure 2Suggested common name: Tung’s Narrow-mouthed FrogHolotype. HC00279 (Figure 2), adult female, collected April 2008 from Gunung Besar Hantu, NegeriSembilan, Peninsular Malaysia (Figure 1) by Orang Asli collectors at an elevation of 800–1400 meters a.s.l.Paratypes. HC00199, male; LSUHC 9166, female, collected from the same location as the holotype onthe same date.FIGURE 2. Holotype of Gastrophrynoides immaculatus (HC00279).Diagnosis. Gastrophrynoides immaculatus is distinguished from all other microhylids by having a long,pointed snout that is 2.6–3.0 times the diameter of the eye; smooth skin; immaculate, grayish brown dorsumwith no white spots; and a single, large, oval, outer metacarpal tubercle.Description of holotype. Adult female, SVL 29.3 mm; habitus robust; head slightly wider than long(HW/HL 1.06), triangular in dorsal profile; snout long, 3.0 times diameter of eye, half the length of the head,rounded in lateral profile, greatly projecting beyond lower jaw, ventral surface of projection concave; canthusrostralis indistinct; lores, flat, sloping; nostrils laterally placed, much closer to end of snout than to eye;internarial width less than interorbital width; interorbital width more than three times width of upper eyelid;eyes small, 25% of head length; pupil rounded, widely dilated; tympanum visible but obscured by skin,subcircular, as wide as high, slightly less than eye diameter; supratympanic fold weak.Skin smooth on all dorsal and ventral surfaces. Forelimbs moderately short, somewhat robust;F3>F2=F4>F1; finger tips expanded into small, round disks bearing circum-marginal grooves; finger disksequal or slightly wider than penultimate phalanges; subarticular and metacarpal tubercles weak; innermetacarpal tubercle small, oval; outer metacarpal tubercle large, oval; fingers free of web. Hind limbsA NEW SPECIES OF MICROHYLIDZootaxa 2124 © 2009 Magnolia Press · 65

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use.Commercial sale or deposition in a public library or website is prohibited.moderate in length, robust;T4>T2=T3=T5>T1; toe tips expanded into small, round disks bearing circum-marginal grooves; toe disksslightly wider than penultimate phalanges; toes free of webbing; inner metatarsal tubercle ellipsoidal ;outermetatarsal tubercle absent (See appendix for additional measurements).Color in life. Uniform grayish brown above; cream with fine brown stippling (only visible undermicroscope) below.Color in alcohol. Dark gray above, dirty white below.Paratypes. Resemble holotype in coloration except the brown stippling beneath the limbs on HC00199and LSUHC 9166 is denser than that of the holotype (See appendix for measurements of type series).Comparisons. Gastrophrynoides immaculatus is distinguished from G. borneensis in having animmaculate, grayish brown dorsum as opposed to a spotted dorsum (Figure 2,3); a longer snout (2.6–3.0 vs.2.0–2.5 times diameter of eye); and a single, large, oval, outer metacarpal tubercle beneath the hand (vs.smaller, paired, rectangular outer metacarpal tubercles in G. borneensis).Distribution. Only known from the type locality in Gunung Besar Hantu, Negeri Sembilan (Figure 1).Natural History. The type series was found between 800–1400 m elevation in water-filled bamboo cutsnot more than 1 m above ground.FIGURE 3. Gastrophrynoides borneensis from Tawau Hills, Sabah. Photograph by Mr. Kueh Boon Hee.TABLE 1. Selected measurements (in mm) of type series. See materials and methods for abbreviations.HC00279 HC00199 LSUHC 9166SVL 29.3 24.4 26.2TiL 12.3 9.4 12.0ED 1.5 1.4 1.6HL 7.9 7.3 7.2HW 8.4 6.1 7.0SL 4.6 4.1 4.2DETN 2.8 2.2 2.1DNTS 1.7 1.4 1.9ELW 1.1 1.0 1.1IOD 3.8 3.1 3.0TD 1.2 1.0 1.4IND 2.7 2.4 2.666 · Zootaxa 2124 © 2009 Magnolia PressCHAN ET AL.

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use.Commercial sale or deposition in a public library or website is prohibited.Etymology. The specific epithet is a latin masculine adjective in reference to the species’ uniformlycolored dorsum, which differentiates it from its only congener, G. borneensis. The proposed common name,Tung’s Narrow-mouthed Frog honors Mr. Leong Sung Tung, who discovered this frog and has contributedimmensely to the study and knowledge of herpetofauna in Peninsular Malaysia.DiscussionIt has been evident for many years that the mountain ranges of Peninsular Malaysia are regions of highherpetological diversity and endemism (e.g., Chan & Grismer 2008; Dring 1979; Grismer et al. 2008, 2009;Grismer 2006, 2007, 2008; Leong & Lim 2003; Wood et al. 2009). However, the majority of upland surveyshave focused only along well known and easily accessible areas (e.g. Genting Highlands, CameronHighlands, Fraser’s Hill, Bukit Larut), which account for only a small fraction of the elevated massifs ofPeninsular Malaysia. The fact that a previously unreported genus has been discovered on an unexploredportion of the Titiwangsa Range brings to attention the dire need for surveys to be conducted in otherunexplored highlands along this extensive mountain corridor. The discovery of a Gastrophrynoides inPeninsular Malaysia is not surprising but is consistent with known prehistoric land connections between theMalay Peninsula, Borneo, Sumatra and Java, which resulted from lowered sea levels during the Pleistoceneglaciations (Inger 1966; Heaney 1986; Hall 1996, 1998). This is reflected in relatively high similarity indicesbetween Borneo and Peninsular Malaysia (Inger & Voris 2001).AcknowledgementsWe thank Robert Inger from the Field Museum of Natural History, Chicago, USA, and Alexander Haas fromUniversity of Hamburg, Germany, for their valuable insights and help in identification. We thank Kueh BoonHee from the Institute for Tropical Biology and Conservation, University Malaysia Sabah for allowing us touse his photograph. For the loan of materials we thank Alan Resetar of the Field Museum of Natural Historyand Aaron M. Bauer for the radiographs of the paratype LSUHC 9166.ReferencesBoulenger, M.A. (1897) Descriptions of new Malay Frogs. Annals and Magazine of Natural History, 19, 106–108.Chan, K.O. & Grismer, L.L. (2008) A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Selangor,Peninsular Malaysia. Zootaxa, 1877, 49–57.De Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: aconceptual unification and terminological recommendation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endlessforms, species and speciation. Oxford University Press, Oxford, pp. 57–75.Dring, J.C.M. (1979) Amphibians and reptiles from northern Terengganu, Malaysia, with descriptions of two newgeckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum of Natural History, 34, 181–241.Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., De Sá, R.O., Channing, A., Wilkinson, M.,Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch,J.D., Green, D.M. & Wheeler, W.C. (2006). The amphibian tree of life. Bulletin of the American Museum of NaturalHistory, 297, 1–370.Grismer, L. L. (1999) An evolutionary classification of reptiles on islands in the Gulf of California, Mexico.Herpetologica, 55, 446–469.Grismer, L. L., Norhayati A., Chan, K. O., Daicus, B., Muin, M. A., Wood, P. L. Jr., Grismer, J. L. (2009) Two newdiminutive species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Peninsular Malaysia. Zootaxa, 2019,40–56.Grismer, L.L. (2006) A new species of Ansonia Stoliczka 1872 (Anura: Bufonidae) from central peninsular Malaysia anda revised taxonomy for Ansonia from the Malay Peninsula. Zootaxa, 1327, 1–21.A NEW SPECIES OF MICROHYLIDZootaxa 2124 © 2009 Magnolia Press · 67

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use.Commercial sale or deposition in a public library or website is prohibited.Grismer, L.L. (2007) A new species of small montane forest floor skink (Genus Sphenomorphus Fitzinger 1843) fromsouthern Peninsular Malaysia. Herpetologica, 63, 544–551.Grismer, L.L. (2008) A new species of insular skink (Genus Sphenomorphus Fitzinger 1843) from the LangkawiArchipelago, Kedah, West Malaysia with the first report of the herpetofauna of Pulau Singa Besar and an updatedchecklist of the herpetofauna of Pulau Langkawi. Zootaxa, 1691, 53–66.Grismer, L.L., Grismer, J.L., Wood, P.L.Jr. & Chan, K.O. (2008) The distribution, taxonomy, and redescription of thegeckos Cnemaspis affinis (Stoliczka 1887) and C. flavolineata (Nicholls 1949) with descriptions of a new montanespecies and two new lowland, karst- dwelling species from Peninsular Malaysia. Zootaxa, 1931, 1–24.Hall, R. (1996) Reconstructing Cenozoic SE Asia. In: Hall, R. & Blundel, D. J. (Eds.), Tectonic evolution of SoutheastAsia. Geological Society Publishing House, Bath, pp. 153–183.Hall, R. (1998) The plate tectonics of Cenozoic SE Asia and thedistribution of land and sea. In: Hall, R. & Holloway, J. D. (Eds.), Biogeography and geological evolution of SoutheastAsia. Brackhuys Publishers, Leiden, pp. 99–132.Heaney, L. R. (1986) Biogeography of small mammals in SE Asia: estimates of rates of colonization, extinction andspeciation. Biological Journal of the Linnean Society, 28, 127–165.Inger, R.F. (1966) The Systematics and Zoogeography of the Amphibia of Borneo. Fieldiana : Zoology. Volume 52. FieldMuseum of Natural History, Chicago, U.S.A. 402 pp.Inger, R. F. & Voris, H. K. (2001) The biogeographical relations of the frogs and snakes of Sundaland. Journal ofBiogeography, 28, 863–891.Inger, R.F. & Stuebing, R.B. (2005) A Field Guide to the Frogs of Borneo. Second Edition. Natural History Publications(Borneo) Sdn Bhd. Kota Kinabalu, Sabah, Malaysia. viii + 201 pp.Kueh, B. H. & Sudin, A. (2008) Gastrophrynoides borneensis range extension. Herpetological Review, 29(3), 363.Leong, T. M. & Lim, B.L. (2003) A new species of Rana (Amphibia: Anura: Ranidae) from the highlands of the MalayPeninsula. Raffles Bulletin of Zoology, 51, 115–122.McGuire, J. A., Brown, R. M., Mumpuni, Riyanto, A. & Andayani, N. (2007) The flying lizards of the Draco lineatusgroup (Squamata: Iguania: Agamidae): A taxonomic revision with descriptions of two new species. HerpetologicalMonographs, 21, 180–213.Van der Meijden, A., Vences, M., Hoegg, S., Boistel, R., Channing, A. & Meyer, A. (2007) Nuclear gene phylogeny ofnarrow-mouthed toads (Family Microhylidae) and a discussion of competing hypotheses concerning theirbiogeographical origins. Molecular Phylogenetics and Evolution, 44, 1017–1030.Wood, P. L. Jr., Grismer, J. L., Grismer, L. L., Norhayati, A., Chan, K. O., Bauer, A. (2009) Two new montane species ofAcanthosaura Gray, 1831 (Squamata: Agamidae) from Peninsular Malaysia. Zootaxa, 2012, 28–46.Appendix. Materials examinedGastrophrynoides borneensisMalaysia: Sarawak: 3 rd Division, Kapit District, Mengiong River, Nanga Tekalit camp (FMNH 145314).Sarawak: 4 th Division, Bintulu District, Tubau camp on Sungai Pesu (FMNH 157179).Sarawak: 7 th Division, Kapit District, Nanga Tekalit (FMNH 223101).68 · Zootaxa 2124 © 2009 Magnolia PressCHAN ET AL.