A Review of the Frog Genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae)
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Zeylanica, ISSN 1391–6270. May, 2001. Vol. 6, No. 1, pp. 1–112, 22 figs., 2 tabs. © 2001, Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka. A review of the frog genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae) Franky Bossuyt* and Alain Dubois** * Free University of Brussels, Biology Deparment, Unit of Ecology and Systematics, Pleinlaan 2, 1050 Brussels, Belgium. ** Laboratoire des Reptiles et Amphibiens, Muséum national d’Histoire naturelle, 25 rue Cuvier, 75005 Paris, France. Abstract This paper is devoted to a review of the specific taxonomy of the frog genus Philautus Gistel, 1848. From 1822 to 1999, 177 nominal species were either described as members of this genus, or of other genera but subsequently referred to this genus. We tried to review the available information on the taxonomic status of these 177 names and the status of their name-bearing types. As a result of this review, 143 types are known to be extant, including 19 lectotypes and 8 neotypes designated and/or described in the present paper. In conclusion of this preliminary analysis, we provisionally distribute these 177 names in 84 valid species names in the genus Philautus, 37 invalid synonyms of the latter names, and 56 nominal species now referred to other genera. These results are highly provisional, both at specific and supra- specific levels. Additional works, using various characters and methods, will be necessary to confirm or reject the validity of a number of these species, and many additional species clearly remain to be discovered and described in the whole range of this genus. At supraspecific level, the taxonomy we use (a single genus Philautus with three subgenera) is also highly provisional, as the generic taxonomy of the whole subfamily Rhacophorinae is in strong need of revision. The present work will provide clear nomenclatural bases for future works on the phylogeny and taxonomy of this difficult group. Contents Abbreviations ........................................................................................................ 2 Introduction ........................................................................................................... 3 Historical summary of the taxonomy of the genus Philautus Gistel, 1848 ..... 3 Familial taxonomy ......................................................................................... 3 Generic taxonomy .......................................................................................... 4 Specific taxonomy .......................................................................................... 7 Provisional supraspecific taxonomy of the genus Philautus Gistel, 1848 ....... 7 Provisional subgenera ................................................................................... 7 Species excluded from the genus Philautus .............................................. 8 Methodology followed in the list of nominal species .................................. 8 Chronological commented list of available and unavailable scientific species-group names for frogs originally referred to the genera Ixalus Duméril & Bibron, 1841 or Philautus Gistel, 1848, and/or subsequently referred to these genera ......................................... 12 Description of type-specimens .......................................................................... 62 List of currently recognized taxa and synonyms for frogs of the genus Philautus Gistel, 1848 ..................................................................................... 84 Conclusion ............................................................................................................. 92 Acknowledgements .............................................................................................. 96 Literature cited ...................................................................................................... 97 Index to scientific names ..................................................................................... 102 BOSSUYT & DUBOIS Abbreviations td1 to td5. Width of disk of toes 1 to 5. TFOL. Length of tarsus and foot (from base of tarsus Measurements to tip of fourth toe). nm. Measurement not taken on this specimen. TFTF. Distance from maximum incurvation of web Body between third and fourth toe to tip of fourth toe, SVL. Snout-vent length. toes being spread. Head TL. Tibia length. EL. Eye length. TW. Maximum tibia width. EN. Distance from anterior corner of eye to nostril. tw1 to tw5. Width of toes 1 to 5. HL. Head length (from posterior corner of mandible to tip of snout). Collection numbers, persons and museums HW. Head width, at the angle of jaws. cnu. Collection number(s) unknown. IBE. Distance between posterior corner of eyes. ex. Original collection or collection number, now IFE. Distance between anterior corner of eyes. changed. IN. Internarial distance. IUE. Minimum distance between upper eyelids. AD. Alain Dubois. MBE. Distance from posterior corner of mandible to FB. Franky Bossuyt. posterior corner of eye. MFE. Distance from posterior corner of mandible to ANSP. Academy of Natural Sciences, Philadelphia, anterior corner of eye. Pennsylvania, USA. MN. Distance from posterior corner of mandible to BMNH. Natural History Museum, London, United nostril. Kingdom. NS. Distance from nostril to tip of snout. BNHS. Bombay Natural History Society, Bombay, SL. Distance from anterior corner of eye to tip of Maharashtra, India. snout. CAS. California Academy of Sciences, San Francisco, TYD. Maximum tympanum diameter. California, USA. TYE. Distance between tympanum and posterior CAS-SU. Stanford University collection, California corner of eye. Academy of Sciences, San Francisco, California, UEW. Maximum width of upper eyelid. USA. Forelimb CCB. Central College, Bangalore, Karnataka, India. fd1 to fd4. Width of disk of fingers 1 to 4. CIB. Chengdu Institute of Biology, Adacemia Sinica, FLL. Forelimb length (from elbow to base of outer Chengdu, Sichuan, China. palmar tubercle). CM. Carnegie Museum, Pittsburgh, Pennsylvania, fw1 to fw4. Width of fingers 1 to 4. USA. HAL. Hand length (from base of outer palmar EHT. Edward H. Taylor collection. tubercle to tip of third finger). FMNH. Field Museum of Natural History, Chicago, TFL. Third finger length (from base of first Illinois, USA. subarticular tubercle). IRSNB. Institut Royal des Sciences Naturelles de Hindlimb Belgique, Bruxelles, Belgium. FFTF. Distance from maximum incurvation of web KIZ. Kunming Institute of Zoology, Academia Sinica, between fourth and fifth toe to tip of fourth toe, Kunming, Yunnan, China. toes being spread. KM. Muzeum Przyrodnicze Uniwersytetu Jagiel- FL. Femur length (from vent to knee). lonskiego, Kraków, Poland. FOL. Foot length (from base of inner metatarsal LZUH. Laboratoire de Zoologie, Université de Hanoi, tubercle to tip of fourth toe). Vietnam. FTL. Fourth toe length (from base of first subarticular MAS. Malcolm A. Smith collection. tubercle). MCZ. Museum of Comparative Zoology, Cambridge, IMT. Length of inner metatarsal tubercle. Massachusetts, USA. ITL. Inner toe length. MNHN. Muséum National d’Histoire Naturelle, Paris, MTFF. Distance from distal edge of metatarsal France. tubercle to maximum incurvation of web MSNG. Museo Civico di Storia Naturale Giacomo Doria, between fourth and fifth toe, toes being spread. Genova, Italy. MTTF. Distance from distal edge of metatarsal tub- NHMB. Naturhistorisches Museum, Basel, Switzerland. ercle to maximum incurvation of web between NMW. Naturhistorisches Museum, Wien, Austria. third and fourth toe, toes being spread. NTUM. National Taiwan University, Taipei, Taiwan. 2 Zeylanica REVIEW OF PHILAUTUS PBS - Philippine Bureau of Science, Philippines. necessary to review the nomenclatural history of all RMNH. Rijksmuseum van Natuurlijke Historie, Leiden, nominal taxa once referred to this genus, in order to Netherlands. provide a sound basis for further morphological, SMF. Forschunsinstitut und Natur-Museum anatomical, morphometric, bioacoustic and molecular Senckenberg, Frankfurt-am-Main, Germany. studies of these frogs. This will be particularly useful in USNM. National Museum of Natural History, Wash- regions where numerous species of this group remain ington, DC, USA. to be described, as seems to be particularly the case in YU. Yunnan University, Kunming, China. Sri Lanka (Pethiyagoda & Manamendra-Arachchi, ZMA. Zoologisch Museum, Universiteit van Amster- 1998). This paper is devoted to the detailed presentation dam, Netherlands. of the results of this work. We stress that this is largely a ZMB. Zoologisches Museum, Berlin, Germany. nomenclatural review of the genus Philautus, not a ZSIC. Zoological Survey of India, Calcutta, West Ben- taxonomic revision, a much-needed work still far from gal, India. being possible for the time being. However, since ZSIM. Zoological Survey of India, Madras, Tamil Nadu, nomenclature is not completely independent of India. taxonomy, we had to address various taxonomic ZSIS. Zoological Survey of India, Shillong, Meghalaya, problems in the course of this nomenclatural review; India. we also provide a few distributional, historical and other pieces of information, when available: these may be useful to future workers on this genus. In order to Introduction stabilize definitely the use of several problematic names or of names based on syntypes, we designate and Anyone who has spent some nights in forested areas of describe below a number of neotypes and lectotypes. South or South-East Asia during the rainy season