Abstract
A new diminutive species of microhylid frog (genus Microhyla) is described from the Matang Range, Sarawak, Malaysian Borneo. The new species is an obligate of the pitcher plant, Nepenthes ampullaria, breeding in senescent or mature pitchers, and is Old World’s smallest frog and one of the world's tiniest: adult males range between SVL 10.6– 12.8 mm (n = 8). The new species is diagnosable from congeners in showing dorsum with low tubercles that are relatively more distinct on flanks; a weak, broken, mid-vertebral ridge, starting from forehead and continuing along body; no dermal fold across forehead; tympanic membrane and tympanic annulus absent; Finger I reduced to a nub proximal to Finger II in males; toe tips weakly dilated; phalanges with longitudinal grooves, forming two scale-like structures; webbing on toe IV basal; toes with narrow dermal fringes; inner and outer metatarsal tubercles present; and dorsum brown with an hour-glass shaped mark on scapular region. Miniaturization and reduced webbing may be the result of navigation on the slippery zone of pitchers, situated below the peristome.References
Alberch, P. & Gale, E.A. (1985) A developmental analysis of an evolutionary trend: digital reduction in amphibians. Evolution, 39, 8¨C23.
Bain, R. & Truong, N.Q. (2004) Three new species of narrow-mouth frogs (genus: Microhyla) from Indochina, with comments on Microhyla annamensis and Microhyla palmipes. Copeia 2004, 507¨C524.
Berry, P.Y. (1975) The Amphibian Fauna of Peninsular Malaysia. Tropical Press, Kuala Lumpur, x + 130 pp.
Biju, S.D., Van Bocxlaer, I., Giri, V.B., Roelants, K., Nagaraju, J. & Bossuyt, F. (2007) A new nightfrog, Nyctibatrachus minimus sp. nov. (Anura: Nyctibatrachidae): the smallest frog from India. Current Science, 93, 854¨C858.
Clarke, C. (1997) Nepenthes of Borneo. Natural History Publications (Borneo) Sdn. Bhd. and Science and Technology Unit, Kota Kinabalu, xii + 207 pp.
Clarke, C. & Lee, C. (2004) A Pocket Guide. Pitcher Plants of Borneo. Natural History Publications (Borneo) Sdn. Bhd. and Science and Technology Unit, Kota Kinabalu, vi + 81 pp.
Das, I., Jankowski, A., Makmor, M.I. & Haas, A. (2007) Species diversity, elevational distribution and reproductive modes in an amphibian community at the Matang Range, Sarawak (Borneo). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 104, 1¨C34.
Das, I. & Leh, C. (2005) A legacy of Wallace: Sarawak Museum and the history of herpetological research in Sarawak. In: A. A. Tuen & Das, I. (Eds). Wallace in Sarawak¨C 150 Years Later: Proceedings of an International Conference on Biogeography and Biodiversity. Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, Kota Samarahan. pp: 57¨C65.
Das, I., Yaakob, N. & Sukumaran, J. (2007) A new species of Microhyla (Anura: Microhylidae) from the Malay Peninsula. Hamadryad, 32, 304¨C314.
Dover, C. (1928) Part III.¨C Batrachia. In: Notes on the fauna of pitcher-plants from Singapore Island. Journal of the Malayan Branch of the Royal Asiatic Society, 6, 13.
Dring, J.C. (1987) Bornean treefrogs of the genus Philautus (Rhacophoridae). Amphibia-Reptilia, 8, 19¨C47.
Duellman, W.E. & Hedges, S.B. (2008) Two new minute species of Phrynopus (Lissamphibia: Anura) from the Cordillera Oriental in Peru. Zootaxa, 1675, 59¨C66.
Dutta, S.K. & Ray, P. (2000) Microhyla sholigari, a new species of microhylid frog (Anura: Microhylidae) from Karnataka, India. Hamadryad, 25, 38¨C44.
Eriksson, T. (2003) AutoDecay. Available from: http://www.bergianska.se/index_forskning_soft.html
Estrada, A.R. & Hedges, S.B. (1996) At the lower size limit in tetrapods: a new diminutive frog from Cuba (Leptodactylidae: Eleutherodactylus). Copeia, 1996, 852¨C859.
Evans, B.J., Brown, R.M., McGuire, J.A., Supriatna, J., Andayani, N., Diesmos, A., Iskandar, D., Melnick, D.J. & Cannatella, D.C. (2003) Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones. Systematic Biology, 52, 794–819.
Fernando, P. & Siriwardhane, M. (1996) Microhyla karunaratnei (Anura: Microhylidae), a new species of frog endemic to Sri Lanka. Journal of South Asian Natural History, 2, 135¨C142.
Frost, D.R. (1985) Amphibian Species of the World. A Taxonomic and Geographic Reference. Allen Press and Association of Systematics Collection, Lawrence, Kansas, 732 pp.
Frost, D.R. (2009) Amphibian Species of the World 5.3, An Online Reference. Available from http://research.amnh.org/vz/herpetology/amphibia/index.php). American Museum of Natural History, New York. (Accessed 21 October 2009)
Frost, D.R., Grant, T., Faivovich, J.N., Bain, R.H., Haas, A., Haddad, C.F.B., De S¨¢, 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) Amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1¨C370.
Gorb, E., Haas, K., Henrich, A., Enders, S., Barbakadse, N. & Gorb, S. (2005) Composite structure of the crystalline epicuticular wax layer of the slippery zone in the pitchers of the carnivorous plant Nepenthes alata and its effect on insect attachment. Journal of Experimental Biology, 208, 451¨C462.
Gosner, K.L. (1960) A simplified table for staging anuran embryos and larvae, with notes on identification. Herpetologica, 16, 183¨C190.
Haas, A. (2003) The phylogeny of frogs as inferred from primarily larval characters (Amphibia: Anura). Cladistics, 19, 23¨C89.
Hedges, S.B. & Thomas, R. (2001) At the lower size limit in amniote vertebrates: a new diminutive lizard from the West Indies. Caribbean Journal of Science, 37, 168¨C173.
Hillis, D.M., Moritz, C. & Mable, B.K. (1996) Molecular Systematics. Second Edition. Sinauer Associates, Sunderland, MA, 655 pp.
Huson, D.H., Richter, D.C., Rausch, C., Dezulian, T., Franz, M. & Rupp, R. (2007) Dendroscope: An interactive viewer for large phylogenetic trees. BMC Bioinformatics, 2007, 8, 460.
Huelsenbeck, J.P. & Ronquist, F.R. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Inger, R.F. (1966) The systematics and zoogeography of the Amphibia of Borneo. Fieldiana Zoology, 52, 1¨C402. Reprinted 1990, Lun Hing Trading Company, Kota Kinabalu.
Inger, R.F. (1989) Four new species of frogs from Borneo. Malayan Nature Journal, 42, 229¨C243.
Inger, R.J. & Frogner, K.J. (1980) New species of narrow-mouth frogs (genus Microhyla) from Borneo. Sarawak Museum Journal, 27, 311¨C324.
Inger, R.F. & Stuebing, R.B. (1997) A Field Guide to the Frogs of Borneo. Natural History Publications (Borneo) Sdn Bhd/Science and Technology Unit, Sabah, Kota Kinabalu, x + 205 pp. Bahasa Malaysia edition 1999, Panduan lapangan katak-katak Borneo. Natural History Publications (Borneo) Sdn Bhd/Jabatan Muzium Sabah, Kota Kinabalu, x + 225 pp.
Iskandar, D.T. & Colijn, E. (2000) Preliminary checklist of southeast Asian and New Guinean herpetofauna. I. Amphibians. Treubia, 31(3)(Supplement), 1¨C133.
Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066.
Kiew, B.-H. (1987) An annotated checklist of the herpetofauna of Ulu Endau, Johore, Malaysia. In: Malaysian Heritage & Scientific Expedition: Endau-Rompin. Malayan Nature Journal, 41, 413¨C423.
Lehr, E. & Catenazzi, A. (2009) A new species of minute Noblella (Anura: Strabomantidae) from southern Peru: the smallest frog of the Andes. Copeia, 2009, 148¨C156.
Lehtinen, R.M., Lanonoo, M.J. & Wassersug, R.J. (2004) Phytotelm-breeding anurans: past, present and future research. In: Lehtinen, R.M. (Ed). Ecology and Evolution of Phytotelm-breeding Anurans. Miscellaneous Publications, Museum of Zoology, University of Michigan 193, 1¨C9.
Leong, T.M. & Chou, L.M. (1999) Larval diversity and development in the Singapore Anura (Amphibia). The Raffles Bulletin of Zoology, 47, 81¨C137.
Leviton, A.E., Anderson, S.C., Gibbs, R.H., Heal, E. & Dawson, C.E. (1985) Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985, 802¨C832.
Lim, K.K.P. & Ng, P.K.L. (1991) Nepenthiphilous larvae and breeding habits of the sticky frog, Kalophrynus pleurostigma Tschudi (Amphibia: Microhylidae). Raffles Bulletin of Zoology, 39, 209¨C214.
Malkmus, R., Manthey, U., Vogel, G., Hoffmann, P. & Kosuch, J. (2002) Amphibians & Reptiles of Mount Kinabalu (North Borneo). Koeltz Scientific Books, Königstein, 424 pp.
Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien S¨¹dostasiens. Natur und Tier Verlag, M¨¹nster, 512 pp.
Matsui, M. (2009) A new species of Philautus (Amphibia, Anura, Rhacophoridae) from the lowland of Sarawak, western Borneo. Zoological Science, 26, 437¨C442.
Matsui, M., Ito, H., Shimada, T., Ota, H., Saidapur, S.K., Khonsue, W., Tanaka-Ueno, T. & Wu, G.-F. (2005) Taxonomic relationships within the Pan-Oriental narrow-mouth toad, Microhyla ornata as revealed by mtDNA analysis (Amphibia, Anura, Microhylidae). Zoological Science, 22, 489¨C495.
Misof, B. & Misof, K. (2009) A Monte Carlo approach successfully identifies randomness of multiple sequence: a more objective approach of data exclusion. Systematic Biology, 58, 21¨C34.
Nieden, F. (1923) Anura I. Subordo Aglossa und Phaneroglossa, Sectio I Arcifera. Das Tierreich, 46, i¨Cxxxii + 1¨C584.
Orton, G.L. (1957) The bearing of larval evolution on some problems in frog classification. Systematic Zoology, 6, 79¨C86.
Parker, H.W. (1926) The brevicipitid frogs of the genus Microhyla. Annals & Magazine of Natural History, Series 10, 2, 473¨C499.
Parker, H.W. (1934) A Monograph of the Frogs of the Family Microhylidae. British Museum (Natural History), London, (1) + viii + 208 pp. Reprinted 1966, Johnson, New York.
Phillipps, A. & Lamb, A. (1988) Pitcher plants of East Malaysia and Brunei. Nature Malaysiana, 13(4), 8¨C27.
Phillipps, A., Lamb, A. & Lee, C.C. (2008) Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu/Royal Botanic Gardens, Kew, viii + 298 pp.
Pillai, R.S. (1977) On two frogs of the family Microhylidae from Andamans including a new species. Proceedings of the Indian Academy of Sciences, 86B, 135¨C138.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Russell, R.W. (2008) Spring peepers and pitcher plants: a case of commensalism? Herpetological Review, 39(2), 154¨C155.
Smith, M.A. (1925) On a collection of reptiles and amphibians from Mt. Murud, Borneo. Journal of the Sarawak Museum, 3, 5¨C14.
Smithe, F.B. (1975) Naturalist¡¯s Color Guide. Parts I and II. American Museum of Natural History, New York, Part I: 8 pp + 18 colour swatches; Part II: xiii + 229 pp.
Smithe, F.B. (1981) Naturalist¡¯s Color Guide. Part III. American Museum of Natural History, New York, (iv) + 37 pp.
Stocsits, R.R., Letsch, H., Hertel, J., Misof, B. & Stadler, P.F. (2009) Accurate and efficient reconstruction of deep phylogenies from structured RNAs. Nucleic Acids Research, 37, 6184–6193.
Swofford, D.L. (2002) PAUP*4 beta. Phylogenetic Analysis Using Parsimony (and Other Methods). Software package. Sinauer Associates, Inc. Publishers, Sunderland, Massachusetts.
Taylor, E.H. (1962) The amphibian fauna of Thailand. University of Kansas Science Bulletin, 63(8), 265¨C599; errata (= 1 p).
van Kampen, P.N. (1923) The Amphibia of the Indo-Australian Archipelago. E. J. Brill, Leiden, xii + 304 pp.
Vences, M. & Glaw, F. (1991) Revision der Gattung Stumpffia Boettger 1881 aus Madagaskar mit Beschreibung von zwei neuen Arten (Amphibia, Anura, Microhylidae). Acta Biologica Benrodis, 3, 203¨C219.
Vences, M., Thomas, M., Bonett, R. & Vieites, D.R. (2005) Deciphering amphibian diversity through DNA barcoding: chances and challenges. Philosophical Transactions of the Royal Society B: Biological Sciences, 360, 1859¨C1868.
Wu, L., Diong, G. & Xu, R.-H. (1987) [Amphibian Fauna of Guizhou.] Guizhou Peoples Press, Guiyang, (7) + 4 + 192 pp, Pls. 46, folding map 1 [in Chinese.]
Yang, D.-T. (Ed.). (1991) [Amphibian Fauna of Yunnan.] China Forestry Publ. House, Beijing, (1) + (1) + (3) + iv + 259 pp. [in Chinese; authors: S.-M. Li, W.-Z. Liu, S.-Q. L¨¹, B.-L. Wu & D.-T. Yang.]
Ye, R.-Q., Fei, L. & Hu, S.-Q. (1986) Early stages of the embryonic development in Bufo melanostictus. Acta Herpetologica Sinica, new series 5(3), 185¨C188; 2 plates (in Chinese.)
Yong, H.-S., Ng, C.K. & Ismail, R. (1988) Conquest of the land: direct development in a Malaysian Philautus tree frog. Nature Malaysiana, 13(4), 4¨C7.