(continued from inside cover) The Raffles Bulletin …An International Journal of Southeast Asian...

An International Journal of Southeast Asian Zoology

Supplement No. 27 22 November 2013

The Raffles Bulletinof Zoology

Published by the Department of Biological Sciences, National University of Singapore

Articles appearing in this journal are indexed in: SCIENCE CITATION INDEX® CURRENT CONTENTS® AGRICULTURE, BIOLOGY & ENVIRONMENTAL SCIENCE; SCISEARCH® RESEARCH ALERT® BIOLOGICAL ABSTRACTS® CAMBRIDGE SCIENTIFIC ABSTRACTS® AQUATIC SCIENCES & FISHERIES ABSTRACTS

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The Fishes of the Inland Waters of Southeast Asia:

A Catalogue and Core Bibliography of the Fishes Known to Occur in Freshwaters, Mangroves and Estuaries

Maurice Kottelat

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SUPPLEMENTS PUBLISHED(continued from inside cover)

7. Allwood, A. J., A. Chinajariyawong, R. A. I. Drew, E. L. Hamacek, D. L. Hancock, C. Hengsawad, J. C. Jipanin, M. Jirasurat, C. Kong Krong, S. Kritsaneepaiboon, C. T. S. Leong & S. Vijaysegaran, 1999. Host plant records for fruit fl ies (Diptera: Tephritidae) in Southeast Asia. Raffl es Bulletin of Zoology, Supplement 7: 1–92.

8. Ng, P. K. L. & K. S. Tan (eds.), 2000. The biodiversity of South China Sea. Raffl es Bulletin of Zoology, Supplement 8: 1–673.

9. Kurahashi, H. & F. R. Magpayo, 2000. Blow fl ies (Insecta: Diptera: Calliphoridae) of the Philippines. Raffl es Bulletin of Zoology, Supplement 9: 1–78.

10. Jefferson, T. A. & B. D. Smith (eds.), 2002. Facultative freshwater cetaceans of Asia: Their ecology and conservation. Raffl es Bulletin of Zoology, Supplement 10: 1–187.

11. Ng, P. K. L., D. Wowor & D. C. J. Yeo (eds.), 2002. Scientifi c results of the Anambas Expedition 2002. Raffl es Bulletin of Zoology, Supplement 11: 1–130.

12. Yeo, D. C. J., P. K. L. Ng & R. Pethiyagoda (eds.), 2005. Contributions to biodiversity exploration and research in Sri Lanka. Raffl es Bulletin of Zoology, Supplement 12: 1–434.

13. Kottelat, M. & D. C. J. Yeo (eds.), 2005. Southeast Asian freshwater fi sh diversity. Raffl es Bulletin of Zoology, Supplement 13: 1–208.

14. Tan, H. & R.-Q. Jan (eds.), 2007. Proceedings of the 7th Indo-Pacifi c Fish Conference. Raffl es Bulletin of Zoology, Supplement 14: 1–434.

15. Wang, L. K. & C. J. Hails, 2007. An annotated checklist of birds of Singapore. Raffl es Bulletin of Zoology, Supplement 15: 1–179.

16. Tan, S. H. & P. K. L. Ng (eds.), 2007. Crustacean Supplement 1. Raffl es Bulletin of Zoology, Supplement 16: 1–357.

17. Ng, P. K. L., D. Guinot & P. J. F. Davie, 2008. Systema Brachyurorum: Part 1. An annotated checklist of extant brachyuran crabs of the world. Raffl es Bulletin of Zoology, Supplement 17: 1–286.

18. Bieler, R., K. Chalermwat, P. M. Mikkelsen, K. S. Tan & E. Wells (eds.), 2008. Molluscs of Eastern Thailand: Proceedings of the International Marine Bivalve Workshop, Chantaburi, Thailand, August–September 2005, with contributions on other molluscan groups. Raffl es Bulletin of Zoology, Supplement 18: 1–264.

19. Tan, S. H. & I.-S. Chen (eds.), 2008. Aquatic biodiversity of the South China Sea. Raffl es Bulletin of Zoology, Supplement 19: 1–292.

20. Tan, S. H. & M. E. Y. Low (eds.), 2009. Crustacean Supplement II. Raffl es Bulletin of Zoology, Supplement 20: 1–307.

21. De Grave, S., N. D. Pentcheff, S. T. Ahyong, T.-Y. Chan, K. A. Crandall, P. C. Dworschak, D. L. Felder, R. M. Feldmann, C. H. J. M. Fransen, L.Y. D. Goulding, R. Lemaitre, M. E. Y. Low, J. W. Martin, P. K. L. Ng, C. E. Schweitzer, S. H. Tan, D. Tshudy & R. Wetzer, 2009. A classifi cation of living and fossil genera of decapod crustaceans. Raffl es Bulletin of Zoology, Supplement 21: 1–109.

22. Tan, K. S. (ed.), 2009. Fourteenth International Marine Biology Workshop 2006: The marine fl ora and fauna of Singapore. Raffl es Bulletin of Zoology, Supplement 22: 1–294.

23. Low, M. E. Y. & S. H. Tan (eds.), 2010. Checklists of anomuran decapod curstaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) and marine lobsters of the world. Raffl es Bulletin of Zoology, Supplement 23: 1–181.

24. Davison, G. W. H. & C. S. W. Chia (eds.), 2011. Proceedings of the fi fth International Hornbill Conference, Singapore, 22–25 March 2009. Raffl es Bulletin of Zoology, Supplement 24: 1–176.

25. Koh, L. P., T. M. Lee & M. L. M. Lim (eds.), 2012. Special Memorial Issue: Navjot S. Sodhi (1962–2011). Raffl es Bulletin of Zoology, Supplement 25: 1–289.

26. Kottelat, M., 2013. Conspectus cobitidum: An inventory of the loaches of the world (Teleostei: Cypriniformes: Cobitoidei). Raffl es Bulletin of Zoology, Supplement 26: 1–199.

INSTRUCTIONS TO AUTHORS

Submission of manuscripts. — All manuscripts are to be submitted via email to [email protected], with a list of at least two potential reviewers. Where possible, authors are encouraged to deposit representative material from papers published in the RBZ with the Zoological Reference Collection (ZRC) of the Raffl es Museum of Biodiversity Research (RMBR), National University of Singapore.

Presentation. — Documents produced with Microsoft® Word (.doc) are preferred. Text must be double-spaced throughout. The title page should contain the full title, author’s name, professional affi liation and email address, and a short running title of not more than 35 characters. All articles should be accompanied by an abstract of not more than 300 words and 4–8 key words. All scientifi c names used or proposed must be in accordance with the 4th Edition of the International Commission of Zoological Nomenclature (1999). Telegraphic style is recommended for descriptions, diagnoses, and keys in taxonomic papers. The holotype must be clearly designated and depositories for all type specimens must be clearly stated, including catalogue numbers if possible. Descriptions of new taxa by one author in a paper under another’s name are strongly discouraged (e.g., Lim, in Tan & Ong, 1986). Synonymies must be cited in the short form (taxon, author, year, page), with full references at the end of the paper.

Literature cited. — All references cited in the text, including taxonomic authorities for any scientifi c names, should be included in the LITERATURE CITED section. References are to be cited in the text by the author’s family name or surname and year of publication, e.g., (King et al., 1975; Nakasone & Agena, 1984; Murphy, 1990). Citations are listed at the end of the manuscript in alphabetical and chronological order. Journal references should include year of publication, title of paper, full title of journal, volume number, and page numbers. Book references should include year of publication, title of book chapter, editor (if any), title of book, publisher, city of publication, and the page numbers of the chapter or the book.

Review. — All manuscripts will be sent to an Associate Editor who will have them reviewed by two referees. The Associate Editor decides on provisional acceptance or rejection based on comments submitted by the referees. After acceptance, the manuscript must be revised and emailed to the Associate Editor for verifi cation.

Page and colour page charges. — There are no page or colour page charges. Authors with papers longer than 20 pages are advised to write to the Managing Editor before submission.

Proofs and reprints. — Proofs in PDF format are emailed to authors for correction and approval, together with the copyright transfer and reprint order forms. Reprint orders are taken with returned proofs. Authors will receive an electronic reprint in PDF format for personal use (note that copyright remains with the publisher). Hard copy reprints cost S$1.00 per page without colour plates, and S$2.00 for a page with colour plates.

Detailed instructions to authors are available at http://rmbr.nus.edu.sg/rbz/author.htm

SUPPLEMENTS PUBLISHED

1. Wee, D. P. C. & P. K. L. Ng, 1995. Swimming crabs of the genera Charybdis De Haan, 1883 and Thalamita Latreille, 1829 (Crustacea: Decapoda: Brachyura: Portunidae) from Peninsular Malaysia and Singapore. Raffl es Bulletin of Zoology, Supplement 1: 1–128.

2. Deeleman-Reinhold, C. L., 1995. The Ochyroceratidae of the Indo-Pacifi c region (Araneae). Raffl es Bulletin of Zoology, Supplement 2: 1–103.

3. Ng, P. K. L. & C. T. N. Chuang, 1996. The Hymenosomatidae (Brachyura) of Southern Asia, with notes on other species. Raffl es Bulletin of Zoology, Supplement 3: 1–82.

4. Morioka, H. & C. M. Yang, 1996. A catalogue of the bird specimens in the Singapore Zoological Reference Collection Part I. Struthioniformes–Charadriiformes. Raffl es Bulletin of Zoology, Supplement 4: 1–141.

5. Kurahashi, H., N. Benjaphong & B. Omar, 1997. Blow fl ies (Insecta: Diptera: Calliphoridae) of Malaysia and Singapore. Raffl es Bulletin of Zoology, Supplement 5: 1–88.

6. Sodhi, N. S., H. S. Yong & P. K. L. Ng (eds.), 1999. The biodiversity of Pulau Tioman, Peninsular Malaysia. Raffl es Bulletin of Zoology, Supplement 6: 1–288.

(continues on back cover)

The Raffl es Bulletin of Zoology | R B Z | rmbr.nus.edu.sg/rbz/The Raffl es Bulletin of Zoology (RBZ) is an online, peer-reviewed journal which publishes high quality papers in Taxonomy, Systematics, Ecology, and Conservation Biology of animals from Southeast Asia and its adjacent areas. The Journal aims to build up quality information on the “animal diversity” of Southeast Asia in particular. Papers from outside the stated geographic range that deal with material deposited in the Zoological Reference Collection (ZRC) of the Raffl es Museum of Biodiversity Research (RMBR), National University of Singapore (NUS) will also be published. Both descriptive and experimental papers will be considered. Single species descriptions and ecosystem studies will be considered for publication. Papers outside the stated policy will be accepted at the discretion of the Editors/Editorial Board.

EDITORIAL BOARDBarry W. Brook (Charles Darwin University, Australia) • Chou Loke Ming (NUS, Singapore) • Indraneil Das (Universiti Malaysia Sarawak, Malaysia) • R. A. I. Drew (Griffi th University, Australia) • Hugh A. Ford (University of New England, Australia) • Fabian Herder (Zoologisches Forschungsmuseum Alexander Koenig, Germany) • Michel Jangoux (University of Brussels, Belgium) • Maurice Kottelat (Cornol, Switzerland) • Damir Kovac (Senckenberg Museum, Germany) • Kelvin K. P. Lim (NUS, Singapore) • John E. Randall (Bernice P. Bishop Museum, USA) • Fred Sheldon (Louisiana State University, USA) • Daniel Simberloff (University of Tennessee, USA) • S. H. Tan (NUS, Singapore) • Y. Tsubaki (Center for Ecological Research, Kyoto University, Japan) • E. O. Wilson (Harvard University, USA) • S. Yamagishi (Kyoto University, Japan) • H. S. Yong (University of Malaya, Malaysia)

MANAGING EDITORIAL BOARDEDITOR-IN-CHIEF: Peter K. L. Ng (NUS, Singapore)MANAGING EDITOR: Tan Heok Hui (NUS, Singapore)ASSOCIATE EDITORS: Kevin Conway (Texas A&M University, USA) • Hwang Wei Song (NUS, Singapore) • Zeehan Jaafar (NUS, Singapore) • Sebastian Klaus (Johan Wolgang Goethe-Universität Biologicum, Germany) • Jeffrey Kwik Teik Beng (NUS, Singapore) • Leong Tzi Ming (NUS, Singapore) • Li Daiqin (NUS, Singapore) • Norman Lim T-Lon (University California, Davis, USA) • Rudolf Meier (NUS, Singapore) • Jose Christopher E. Mendoza (NUS, Singapore) • Tohru Naruse (University of Ryukyus, Japan) • James Reimer (University of Ryukyu, Japan) • Frank E. Rheindt (NUS, Singapore) • Tan Koh Siang (TMSI, Singapore) • Tan Siong Kiat (NUS, Singapore) • Peter A. Todd (NUS, Singapore) • Tran Anh Duc (Hanoi University of Science, Vietnam) • Darren C. J. Yeo (NUS, Singapore)SENIOR COPY EDITOR: Hazelina H. T. Yeo (NUS, Singapore)COPY EDITOR: Jeremy W. L. Yeo (NUS, Singapore)PRODUCTION EDITOR: Hazelina H. T. Yeo (NUS, Singapore)EDITORIAL ADMINISTRATOR: Greasi Simon (NUS, Singapore)WEBMASTER: Chua Keng Soon (NUS, Singapore)

PUBLICATION DETAILSThe Raffl es Bulletin of Zoology will consist of a single volume (two issues) each year, continuing the sequence of its two predecessors, Bulletin of the Raffl es Museum (1928–1960) and Bulletin of the National Museum of Singapore (1961–1970). A separately numbered supplement series will be published as and when manuscripts and funding permit. About 14 hard-copies of the Journal will be deposited in major publicly accessible libraries to satisfy Article 8.6 of the Fourth Edition of the International Code of Zoological Nomenclature (1999) so that all new names published in the RBZ are considered to be published and available.

COPYRIGHT AND EXCHANGESAll articles published by the RBZ may be downloaded from http://rmbr.nus.edu.sg/rbz/biblio/ for research purposes. A condition of publication is that authors assign publication rights to The Raffl es Bulletin of Zoology. After publication, authors may use the article without prior permission from the Journal provided acknowledgement is given to the Journal as the original source of publication. It is the responsibility of authors to obtain permission to use copyright material from other sources. For permissions and copyright matters, please contact the Managing Editor at [email protected]. For queries on the purchase of back issues or journal exchanges, please contact the Editorial Administrator at [email protected].

PUBLISHER’S ADDRESSThe Raffl es Bulletin of Zoology • Raffl es Museum of Biodiversity Research • Department of Biological Sciences • National University of Singapore • Block S6 : Level 3 • Science Drive 2 • Singapore 117546 • Republic of SingaporeThe Raffl es Bulletin of Zoology is online at http://rmbr.nus.edu.sg/rbz/ ISSN 0217-2445ISBN 978-2-8399-1344-7

THE RAFFLES BULLETIN OF ZOOLOGY 2013

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THE RAFFLES BULLETIN OF ZOOLOGY 2013 Supplement No. 27: 1–663 http://zoobank.org/urn:lsid:zoobank.org:pub:0B66AE04-C644-43CD-9B76-043848FAA9FE Date of Publication: 22 Nov.2013© National University of Singapore and the author

THE FISHES OF THE INLAND WATERS OF SOUTHEAST ASIA :A CATALOGUE AND CORE BIBLIOGRAPHY OF

THE FISHES KNOWN TO OCCUR IN FRESHWATERS,MANGROVES AND ESTUARIES

Maurice KottelatCase postale 57, CH-2952 Cornol, Switzerland (address for correspondence), and

Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore,6 Science Drive 2, Singapore 117546, Republic of Singapore

Email: [email protected]

CONTENTS

Abstract ......................................................................... 3Introduction ................................................................... 4Basic principles of nomenclature ................................ 10Abbreviations used ...................................................... 15

Class ChondrichthyesSubclass Elasmobranchii

Division SelachiiOrder Orectolobiformes

Family Hemiscylliidae .......................... 18Family Stegostomatidae ........................ 19

Order CarcharhiniformesFamily Carcharhinidae ......................... 20

Division BatoideaOrder Pristiformes

Family Pristidae .................................... 23Order Rajiformes

Family Rhinobatidae ............................. 24Order Myliobatiformes

Family Dasyatidae ................................ 25Family Potamotrygonidae ..................... 28Family Myliobatididae .......................... 29

Class ActinopterygiiDivision Teleostei

Order OsteoglossiformesFamily Osteoglossidae ............................... 30Family Arapaimidae ................................... 31Family Notopteridae ................................... 31

Order ElopiformesFamily Elopidae ......................................... 33Family Megalopidae ................................... 33

Order AlbuliformesFamily Albulidae ........................................ 34

Order AnguilliformesFamily Anguillidae ..................................... 37Family Moringuidae ................................... 39Family Muraenidae .................................... 41Family Ophichthidae .................................. 43Family Muraenesocidae ............................. 48

Order ClupeiformesFamily Pristigastridae ................................. 49Family Engraulididae ................................. 52Family Chirocentridae ................................ 57Family Clupeidae ....................................... 58Family Sundasalangidae ............................. 64

Order GonorhynchiformesFamily Chanidae ......................................... 64

Order CypriniformesFamily Cyprinidae ...................................... 65Family Psilorhynchidae ............................ 171Family Gyrinocheilidae ............................ 173Family Botiidae ........................................ 173Family Vaillantellidae ............................... 176Family Cobitidae ...................................... 176Family Ellopostomatidae .......................... 184Family Barbuccidae .................................. 185Family Balitoridae .................................... 185Family Gastromyzontidae ........................ 191Family Serpenticobitidae ......................... 197Family Nemacheilidae .............................. 198

Order CharaciformesFamily Characidae .................................... 215Family Serrasalmidae ............................... 215

Order SiluriformesFamily Loricariidae .................................. 215Family Amblycipitidae ............................. 216

Kottelat: Inland fishes of Southeast Asia

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Family Akysidae ....................................... 217Family Sisoridae ....................................... 221Family Cranoglanididae ........................... 232Family Siluridae ....................................... 232Family Chacidae ....................................... 240Family Plotosidae ..................................... 240Family Clariidae ....................................... 242Family Ariidae .......................................... 245Family Schilbeidae ................................... 251Family Pangasiidae .................................. 253Family Bagridae ....................................... 256

Order OsmeriformesFamily Plecoglossidae .............................. 268Family Salangidae .................................... 268

Order SalmoniformesFamily Salmonidae ................................... 270

Order AulopiformesFamily Synodontidae................................ 270

Order GadiformesFamily Bregmacerotidae .......................... 271

Order OphidiiformesFamily Carapidae ..................................... 271Family Bythitidae ..................................... 272

Order BatrachoidiformesFamily Batrachoididae ............................. 272

Order LophiiformesFamily Antennariidae ............................... 274

Order MugiliformesFamily Mugilidae ..................................... 275

Order AtheriniformesFamily Telmatherinidae ............................ 283Family Phallostethidae ............................. 284Family Atherinidae ................................... 286

Order BeloniformesFamily Adrianichthyidae .......................... 288Family Hemiramphidae ............................ 290Family Zenarchopteridae ......................... 293Family Belonidae ..................................... 297

Order CyprinodontiformesFamily Aplocheilidae ............................... 299Family Poeciliidae .................................... 300

Order BeryciformesFamily Holocentridae ............................... 301

Order GasterosteiformesFamily Indostomidae ................................ 301Family Syngnathidae ................................ 301

Order SynbranchiformesFamily Synbranchidae .............................. 307Family Chaudhuriidae .............................. 309Family Mastacembelidae .......................... 310

Order ScorpaeniformesFamily Scorpaenidae ................................ 314Family Tetrarogidae ................................. 315Family Synanceidae ................................. 316Family Platycephalidae ............................ 316

Order PerciformesSuborder Percoidei

Family Ambassidae ............................. 318Family Latidae .................................... 323Family Lateolabracidae ...................... 324

Family Percichthyidae ........................ 324Family Serranidae ............................... 325Family Pseudochromidae ................... 328Family Opistognathidae ...................... 329Family Centrarchidae .......................... 329Family Apogonidae ............................. 329Family Sillaginidae ............................. 330Family Carangidae .............................. 331Family Leiognathidae ......................... 337Family Lutjanidae ............................... 340Family Datnioididae ........................... 344Family Lobotidae ................................ 344Family Gerreidae ................................ 345Family Haemulidae ............................. 347Family Nemipteridae .......................... 349Family Lethrinidae .............................. 350Family Sparidae .................................. 352Family Polynemidae ........................... 353Family Sciaenidae ............................... 356Family Mullidae .................................. 360Family Monodactylidae ...................... 362Family Toxotidae ................................ 362Family Drepaneidae ............................ 365Family Chaetodontidae ....................... 365Family Nandidae ................................. 366Family Pristolepididae ........................ 366Family Badidae ................................... 367Family Terapontidae ........................... 368Family Kuhliidae ................................ 371

Suborder LabroideiFamily Cichlidae ................................. 373Family Pomacentridae ........................ 375Family Labridae .................................. 379Family Scaridae .................................. 380

Suborder TrachinoideiFamily Trichonotidae .......................... 382

Suborder BlennioideiFamily Blenniidae ............................... 382

Suborder CallionymoideiFamily Callionymidae ........................ 385

Suborder GobioideiFamily Rhyacichthyidae ..................... 386Family Odontobutidae ........................ 387Family Eleotrididae ............................ 388Family Kraemeriidae .......................... 396Family Gobiidae ................................. 396Family Amblyopidae .......................... 432Family Ptereleotrididae ....................... 435

Suborder KurtoideiFamily Kurtidae .................................. 436

Suborder AcanthuroideiFamily Ephippidae .............................. 437Family Scatophagidae ......................... 438Family Siganidae ................................ 439Family Acanthuridae ........................... 441

Suborder ScombroideiFamily Sphyraenidae .......................... 444Family Scombridae ............................. 446

Suborder AnabantoideiFamily Anabantidae ............................ 446


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Family Helostomatidae ....................... 447Family Osphronemidae ....................... 448

Suborder ChannoideiFamily Channidae ............................... 458

Order PleuronectiformesFamily Paralichthyidae ............................. 462Family Tephrinectidae .............................. 463Family Soleidae ........................................ 463Family Cynoglossidae .............................. 466

Order TetraodontiformesFamily Triacanthidae ................................ 468

ABSTRACT. — There are 3108 valid and named native fish species in the inland waters ofSoutheast Asia between the Irrawaddy and Red River drainages, the small coastal drainagesbetween the Red River and Hainan, the whole Indochinese Peninsula, Andaman and NicobarIslands, Indonesia (excluding Papua Province, Waigeo, Aru [but Kai is included]), and thePhilippines. They belong to 137 families. Their taxonomy and nomenclature are reviewed. Theoriginal descriptions of all 7047 recorded species-group names and 1980 genus-group nameshave been checked in the original works for correct spelling, types, type locality and bibliographicreferences. The bibliography includes about 4700 titles. Synonymies are given, based onpublished information as well as unpublished observations.

The names of 49 introduced species and 347 extralimital taxa cited in the discussions havealso been checked. The original descriptions of all species not present in the covered area butcited as type species of genera have been checked for availability, authorship, date and correctspelling. The availability of some family-group names has been checked when there was suspicionof possible nomenclatural problems.

Bibliographic notes include new informations on the dates of publication of works by, amongothers, Bleeker, Bloch, Heckel and Steindachner and discussion of authorship of names in variousworks.

The main nomenclatural acts are listed below:– type species designation for: Crayracion, Eleotris Scopoli, Eleotris Walbaum, Encheliopus

Cloquet, Gymnorhinus, Oonidus, Pristipoma Cuvier, Sargus Gronow;– type species fixation under Code art. 70.3.1 for: Bdellorhynchus, Desmoprenes, Innoculus,

Paraprotosalanx;– type species fixation under Code art. 70.3.2 for: Centrurophis, Ovoides Duméril, Pseudoscarus,

Rabula, Waitea;– lectotype designation for: Alausa argyrochloris, Atherina endrachtensis, Barbus gardonides,

Barbus lateristriga, Betta patoti, Betta rubra, Carcharhinus commersonii, Clupea cyprinoides,Clupea gigantea, Clupea thrissoides, Crossochilus benasi, Cyprinus clupeoides, Cyprinuslamta, Engraulis rhinorhynchos, Esox alepidotus, Esox argenteus Gmelin, Esox argenteusSchneider, Equula longispinis, Gobiomoroides piso, Gobius niger, Gonorhynchus bimaculatus,Hemiramphus buffonis, Hemiramphus brevirostris, Hemiramphus georgii, Hemiramphusrusselli Valenciennes, Johnius cataleus, Lobotes auctorum, Neostethus borneensis,Parosphromenus parvulus Foersch & Korthaus, Pellona leschenaulti, Raja edentula, Rajaguttata Shaw, Raja narinari, Rasbora trilineata, Synaptura achira, Teuthis brevirostris, Teuthisjavus;

– neotype designation for: Engraulis dussumieri, Lutjanus gymnocephalus, Ovoides fasciatus,Ovum commersoni, Platygaster megalopterus, Scarus schlosseri, Sciaena jaculatrix, Sciaenapentadactyla;

– declaration as nomina protecta: Albula Scopoli, Aplocheilus, Clupea quadrimaculata, Cyprinusbola, Hemiramphus georgii, Hippocampus Rafinesque-Schmaltz, Kuhlia, Lateolabrax,Mastacembelus erythrotaenia, Oligolepis, Pelates, Phyllopteryx, Platycephalus japonicus,Puntius proctozysron, Raja uarnak Gmelin, Selaroides, Toxotes microlepis Günther;

– declaration as nomina oblita: Albula Osbeck, Barbus carassioides, Centranodon japonicus,Clupea mauritiana, Conorynchus, Cyprinus goha, Gobileptes, Hemiramphus brevirostris,Hemiramphus russellii van Hasselt, Hippocampus Perry, Leptaspis, Mastacembelus catenatus,Odontopsis, Percalabrax, Platerome, Platysoma, Pristipoma Quoy & Gaimard, Rajaommescherit, Raja scherit, Raja schoukie, Sphyraena japonica Bloch, Toxotes microlepisBlyth;

Family Monacanthidae ............................. 469Family Tetraodontidae .............................. 469Family Molidae ........................................ 481

Appendices ................................................................ 481Acknowledgements ................................................... 484

BibliographyBibliographic notes .............................................. 485Literature cited ..................................................... 505


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– first reviser action on correct spelling of Barbus platysoma, Bathygobius variabilis,Boleophthalmus novaeguineae, Euchiloglanis dorsoarcus, Gazza equulaeformis, Kurtus,Leiocassis longispinalis, Neocorassius, Pareuchiloglanis namdeensis, Raja uarnak Walbaum;

– first reviser action on precedence of simultaneous publication of original descriptions of:Parosphromenus parvulus, Puntius roloffi;

– first reviser action on precedence of simultaneous synonyms: Chanodichthys over Pseudoculter,Thrissina over Xenengraulis and Scutengraulis, Xenengraulis over Scutengraulis, Apistuslongispinis over A. bougainvillii, Barbus balleroides over B. hypsylonotus, Carassioidesmacropterus over C. argentea, Cyprinus jogia over C. sutiha, Cyprinus pausius over C. musiha,Eleotris ophicephalus over E. madagascariensis, Gobius caninus over G. quadriporus,Macropodus yeni over M. nigrocorpus, Mystus pahangensis over M. johorensis, Placocheilusbibarbatus over P. imbarbatus, Raja mula over R. tajara, Rohita vittata over R. rostellatus,Tetraodon caria over T. gularis;

– Chaetodon macrolepidotus Linnaeus, 1758 (now in Heniochus) has precedence over C.acuminatus Linnaeus, 1758 as ruled by ICZN, 1912 [Opinion 40]. This Opinion has beengenerally ignored.

– Oxygastri of Bleeker (1860c) is not available because it is a descriptive term, and not based onthe genus name Oxygaster;

– new genera: Desmopuntius (type species: Barbus hexazona Weber & de Beaufort, 1912),Oliotius (type species: Capoeta oligolepis Bleeker, 1853), Puntigrus (type species: Barbuspartipentazona Fowler, 1934), Striuntius (type species: Barbus lineatus Duncker, 1904), Pao(type species: Tetraodon leiurus Bleeker, 1850).

The main unsolved nomenclatural problems are:– the type species of Acanthurus is Naso unicornis and an application to ICZN is needed to

retain the name for species currently called Acanthurus;– the status of Siganus and Teuthis awaits a ruling by ICZN.

KEY WORDS. — freshwater fish, brackish water, mangrove, estuaries, taxonomy, nomenclature,Southeast Asia, Singapore, Indonesia, Malaysia, Thailand, Cambodia, Laos, Vietnam, Myanmar,Manipur, Borneo, Sumatra, Java, Sulawesi, Maluku, Palawan, Sundaland, Indochina, Mekong,Red River, Chao Phraya, Salween, Irrawaddy


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Nowadays it seems fashionable among many zoologists, botanists, anatomistsand physiologists to slightly look down upon this kind of systematic research.In our opinion quite unjustly. Though formerly this kind of investigation may

have been overestimated, at the moment one should not relapse into the oppositeerror. Most likely we are now more than ever in urgent need of accurate

descriptions of species […].

P. Harting, 1878, Pieter Bleeker'’s obituary

human eyes. To these species I have not given the benefit ofthe doubt. The now fashionable discussions about crypticspecies do not change the situation: one nucleotide does notmake a species, be it ever so cryptic. Further, in fishes, thecryptic species discovered by molecular techniques that Ihave been told about have been cryptic not because taxono-mists could not distinguish them, but because no trained tax-onomist ever had an opportunity to examine them.

Another limitation of the precautionary approach in South-east Asian fishes is shown by the huge number of 'new' fishspecies that have been described from Vietnam in recentyears. Their description is of a quality that makes it simplyimpossible to even guess whether or not they might be val-id. The identity and possible distinctness of most will re-main in limbo as long as they are not competently re-de-scribed or evaluated.

Similarly a number of families recognised in recent timesare not recognised. Cladistic molecular phylogeny (whichuses principles and mathematical algorithms that were calledphenetics 30 years ago, an approach rejected by cladistics)has the great particularism of creating fluctuating and tran-sient phylogenies. There is even a case of co-authors pub-lishing contradictory phylogenies simultaneously in twopapers (Mayden & Chen, 2010; Tang et al., 2010; see Britz& Conway, 2011a–b). This shows that it is imprudent toinstantly adopt the latest theory and that naming every littletemporary lineage uncovered by molecular analysis has lit-tle justification.

For this catalogue I have examined personally the originaldescriptions of all the species and genera recorded in theinland waters of Southeast Asia, all their synonyms (mak-ing a total of about 7047 nominal species and about 1980nominal genera). I also examined the original descriptionsof the type species of all genera if they were not known inthe area, and those of the 347 taxa cited in the Taxonomicand Nomenclatural Notes but not present in area. All citednomenclatural acts were checked. The availability of all non-fish names cited as senior synonyms of fish genera waschecked. Synonyms based on fossil taxa and which havenever been used for recent taxa are not included.

INTRODUCTION

The present catalogue aims to present the state of the art ofour knowledge of the diversity of freshwater fishes of South-east Asia. Work began in 1986 when I compiled a list of thefreshwater fishes of the Indochinese region (Kottelat, 1989).The list expanded when I worked on a book on the fishes ofwestern Indonesia (Kottelat et al., 1993; Kottelat & Whit-ten, 1996). Initially it was intended to include only the fresh-water species, but the work for the Indonesian fish bookrequired the inclusion of all species that had been recordedin inland waters, that is, including estuaries, most mangroves,etc. In September 2013, the list includes 3107 valid nativespecies, in 707 valid genera and 137 families. Only namedspecies are included. I am aware of about 300 species to benamed soon or already on museum shelves and a fair num-ber of synonyms to re-validate. I expect an additional 500species still awaiting discovery in the wild. In addition, thereare 49 introduced and established species.

This catalogue is not the ultimate inventory of the fishes ofSoutheast Asian inland waters. Many discoveries are stillahead of us and a great amount of work remains to be donebefore we reach an acceptable level of knowledge.

Taxonomy and systematics have two main goals. One is pri-marily of academic interest: the study of the diversity ofliving organisms and their phylogenetic relationships. Theother is of immediate practical interest: inventories, surveys,documentation of biodiversity, and the compilation of iden-tification tools. For the proper management of natural re-sources, we need information on numbers of species andtheir identification now, not sometime in the distant future.If definitive conclusions are not possible with the availabledata, then tentative decisions are needed. As for other com-ponents of environmental management strategies, the pre-cautionary approach should be the rule. In the present con-text, in case of doubt on the distinctness of two species, theprecautionary approach would be to retain them as distinctawaiting (possible) further research.

This precautionary approach, however, has its limits. Thedevelopment of molecular techniques has led some to rec-ognise as 'species' populations distinguished only by a fewnucleotides; complex statistics have been used to justify therecognition of 'species' otherwise not distinguishable by


6

About 6000 publications (in 22 languages) have been ex-amined and about 4700 relevant ones are listed. All citedreferences have been checked in the original publications.The references or the details that I could not check (for ex-ample, a stolen plate in the only copy that I could access ofan antique book) are marked in bold face (3 out of 4700titles). A few references have been checked by trusted col-leagues.

After completion of the catalogue, all names have been cross-checked against other databases. Especially, all names werecross-checked against Eschmeyer's (2013) Catalog of Fish-es (CoF) in 2001. For each entry in CoF disagreeing withmy data, the data were verified again in the original descrip-tions and literature. In 2010-2011 all names were cross-checked one more time, and in cases of disagreement theoriginal literature was checked for a third time. Out of theabout 9785 checked names, the data (spelling, author, date,types, type locality, etc.) for about 3440 differs from thosein CoF (35 %). Most of the differences are minor and oflittle or no nomenclatural consequences (mainly related withtype localities), but a significant number of differences areserious. The problems are more frequent with the pre-1860non-English literature (type series, type localities, type-spe-cies fixations, dates). In these times of on-demand biodi-versity informatics there are too many assumptions madeabout the quality of the data and there seem to have beenfew or no efforts to carefully evaluate the contents of suchlarge databases. An analysis of the types of differences anderrors is in preparation.

A significant number of nomenclatural problems were dis-covered and the application of the International Code ofZoological Nomenclature (hereunder Code) results in a num-ber of nomenclatural changes. In a few cases of changesaffecting family-group names or widely used names, requestshave been presented to the International Commission onZoological Nomenclature (ICZN) to retain these names intheir current usage (Scatophagidae, Ephippidae, Kottelat,2010b; Siganidae, Kottelat, 2013b; Mystus, Kottelat & Ng,2007). In other cases, I considered that the name changesare minor and I simply applied the Code. At the genus level,I consider that changes resulting from the application of theCode do not create more problems than do changes result-ing from the normal increase of our taxonomic knowledgeby the discovery of new taxa, new characters, etc. I consid-er it appropriate to ask the ICZN to retain the current usagein cases of potential confusion resulting from the discoveryof overlooked type species designations. But I consider itunjustified to ask for the suppression of names (usually se-nior homonyms and synonyms); to me, the argument of sup-pressing names for the sake of stability of nomenclature infavour of reputedly 'well-known' names does not hold in ageographic area where new discoveries still abound andwhere the taxonomic system is still very unstable. Underthat logic, dozens of names should be suppressed and thiswould affect the stability of nomenclature by making thepurpose of the Code irrelevant. Also, writing applicationsfor all these minor cases would mean more applications thanthe ICZN receives in a year.

Limitations. — One of the limitations of this catalogue isthat I started it in 1986; the work spanned 24 years and fouroperating systems and unavoidably this caused slight inter-nal inconsistencies in formatting. It has been updated con-tinuously so that the technical content is not affected by this'formatting evolution'. Further, between 1992 and today, twodifferent editions of the International Code of ZoologicalNomenclature (ICZN, 1985, 1999) have been in use, whichdiffer slightly. I have tried to update all entries affected bythe changes but I may have missed some.

Another limitation is that the core target of this work andmy own experience is the 'real' freshwater fishes. My treat-ment of these taxa is probably close to complete. But I amlikely to have missed some records of estuarine species, orsome literature. When I encountered nomenclatural prob-lems concerning freshwater fishes I had no hesitation in tak-ing the necessary actions to clear the problems. When it cameto the same situation with estuarine taxa, I tried to solve theroutine problems but decided not to address the more com-plex ones. However, I discuss these cases and their possiblesolutions where pertinent. The number of marine taxa withnomenclatural problems was unexpectedly high, and manywell and long known genus and family names are involved.

A potential for small errors arose late in the preparation ofthis catalogue. I had long tried to confirm or revise the chro-nology of the many papers published by Pieter Bleeker andit is only late that I obtained the data to establish the se-quence of publication of some 270 papers he published dur-ing his stay in Java (Kottelat, 2011a) and of the Atlas ich-thyologique (Kottelat, 2013c). A number of the names cre-ated by Bleeker appeared more or less simultaneously indifferent papers and journals. A consequence of this revisedchronology is that the now-established dates of availabilityof many names differ from those commonly recognised, andthis has changed the precedence of the different descriptionof a few species, which now may have a different type se-ries, or of some new genera, which now may have differentoriginally included species, thus potentially invalidatingearlier type species designations. I have tried to eliminatethis risk but I expect that some details would have escaped me.

Geographic and habitat coverages. — The geographiccoverage includes all inland water bodies of Southeast Asiabetween (and including) the Kaladan, Irrawaddy and the RedRiver drainages, the small coastal drainages between the RedRiver and Hainan (included), the whole Indochinese Penin-sula, Andaman and Nicobar Islands, Indonesia (excludingPapua Province, Waigeo and Aru [but Kai is included]), andthe Philippines (Fig. 1).

All freshwater species are included. Introduced species thatbecame established are listed (marked by asterisks, *), butwithout complete synonymies; only species that have es-tablished reproducing populations are listed. Species in-habiting the estuaries, brackish lower stetches of rivers,mangroves, etc. are also included. Species occasionally re-ported in freswaters are recorded, although some of therecords or identifications need critical reevaluations (which


7

was beyond the goals of this work). I preferred to be tooinclusive than too exclusive. For these species too, synony-mies are complete and include the nominal species describedfrom freshwater as well as the marine ones; similarly, thegeneric synonymies applying to these genera are complete.Through a lack of familiarity with some groups I may haveoverlooked some synonyms.

Listed names. — All known names are listed, includinginfrasubspecific names, which are given in their originalform. The only names that are not included are those sub-specific or infrasubspecific epithets typicus when they aremerely intended to denote the nominotypical subspecies;such names are usually not nomenclaturally available andshould not be used.

Spellings. — The headings of all generic and specific ac-counts have the correct spelling of all valid names. In thesynonymies, however, all names are given with their origi-nal combination (except that interpolated subgeneric namesare omitted) and with their original spelling, including mis-spellings, capitalised letters, and diacritic marks [ü, è, ñ, etc.].Capitalised letters and diacritic marks are not permitted bythe Code (arts. 27, 28, Glossary) and must be corrected. In-correct original spellings are used only in the synonymies

but they have been corrected in all other circumstances, es-pecially in the discussions under Nomenclatural notes.

Families. — Families are listed following the sequence inNelson (2006), except within Cypriniformes, for whichI follow Šlechtová et al. (2007) and my personal experience.When there is disagreement between authors with regard tothe limits of families or higher categories, I generally fol-lowed common practice, but have noted alternatives.

With a few exceptions I have not searched the synonymiesof family-group names. Note that a family-group name keepsits original author and date even if used at different ranks.For example Leuciscini Bonaparte, 1835 retains Bonaparte,1835 as author, even if treated as subfamily Leuciscinae orfamily Leuciscidae.

Genera and species. — Genera are listed in alphabeticalsequence within family. Species are listed in alphabeticalsequence within genera.

Entries for genera include the valid name of the genus (inbold, as a heading), the name of the genus with the spellingin the original description, the author, the year of publica-tion, the number of the page with principal information. This

Fig. 1. The geographic area covered by the catalogue.


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is followed by information on possible subgeneric status inthe original description, type species, mode of designation,information on possible nomenclatural acts associated withthe name, and grammatical gender. This information is pro-vided for all names considered to be synonyms, in chrono-logical sequence.

Entries for species include the valid name of the species (inbold, as the heading), the name of the species as spelt in theoriginal description, the author, the year of publication, thenumber of the page on which the actual description starts(or where the elements necessary to make the name avail-able occur) and the number of the main illustrations (ignor-ing those showing maps, anatomical details, portrait of col-lector, etc.). This is followed by a block in parentheses withinformation on type locality and primary types, and infor-mation on possible nomenclatural acts associated with thename. If the name is based largely or totally on references tothe older literature, this information is listed first in the pa-renthesed block). This information is provided for all namesconsidered to be synonyms, in chronological sequence.

Additional information, if needed, is listed under Nomen-clatural notes and Taxonomic notes. When names are citedunder Notes, which are not mentioned elsewhere in the text,I usually (but not always) added the same data for that nameat the end of the paragraph, in brackets.

Transliteration of non-Latin alphabets. — Author names,place names, and journal names in non-Latin alphabets, andin languages using other notations, have been transcribed;titles of books and papers have been translated. When a tran-scription is used in the original work (e.g. in the text, in anabstract, in a table of contents), the same spelling is usedhere. There are some inconsistencies as it happens that tran-scriptions or translations used in abstracts or tables of con-tents may be different from the actual title of a paper. Fre-quently, transcription systems have changed with time andno standardisation has been attempted here. Older bibliog-raphies or indexes may have used earlier transcription sys-tems and I consider that a standardised use could actuallycomplicate bibliographic search, especially for those notfamiliar with these languages.

Unfortunately, some accents and diacritic marks may havedisappeared as standard western European keyboards andsoftware do not support them. This especially applies in thecase of the Vietnamese alphabet.

Type localities. — The type locality is the locality at whichthe holotype, lectotype or neotype was collected. Althoughmentioned in the Code, the type locality has no nomencla-tural role. Simply, it is a convenient wording, it is shorter tosay type locality than 'the locality at which the primary name-bearing type was collected', or to give the locality data infull.

In case there is no primary type but a series of syntypesfrom different localities, the type locality is the sum of allthe localities of the syntypes, and all their localities are list-

ed (separated by a slash [ / ] where clarity requires it). Invery few cases (when the list of localities of syntypes isvery extensive), I have merely given a general descriptionof the localities. Localities are usually given with the origi-nal spelling; this sometimes results in different spellingsbeing used for the same locality under different headings;I have tried to introduce some consistency, but only in caseswhere I was certain that the different spellings were reallyreferring to the same place, or when the different spellingswere used for the locality of the very same specimen, orreferring to the very same bibliographic source. Alternativespellings or modern equivalents are given in square brack-ets, but this has not been systematically attempted. Localitydescriptions have been translated into English when possi-ble and/or justified; in some cases, words meaning river,lake, etc. are part of the name in the original language andthey have not been deleted in order to avoid ambiguitieswhen using local maps [but the word river, lake, etc. hasbeen added]. Local names have been used, except for a fewwell known rivers and lakes with a common English nameused in international literature (e.g. Mekong, Irrawaddy,Salween, Ganges, Red River). For most localities, when fea-sible I have tried to add information on present politicalentities (country, province, state, etc.) and river basin as anaid to the reader. For larger topographic features that haveseveral different names, a single one has been consistentlyused; this especially applies to those traversing differentcountries (e.g. Mekong River); I usually retained the nameeasiest to find for readers not familiar with local toponymyor, when it exists, the English name used in internationalliterature (e.g. Salween River and not Nu Jiang, Salawin,Thanlwin, Salouen, or fGyl mo rNGul chu [a transcriptionfrom Tibetan language]; Irrawaddy and not Ayeyarwady;Red River and not Song Hong, Yuan Jiang or Fleuve Rouge).

As the work on the check-list spanned more than 24 years,it is likely that some of the earlier entries might be in a slightlydifferent format than the latest ones.

Infrasubspecific names and nomina nuda having no nomen-clatural status, they do not have type specimens and there-fore do not have a type locality and I thus list only a 'local-ity', when justified. Localities are usually not indicated forinfrasubspecific names based on aberrant specimens; theyare given only if the name has been created for a particulargeographical form.

When a neotype has been designated, the type locality is thelocality of the neotype. The original type locality [the local-ity of the primary type mentioned in the original descrip-tion], if different, is usually listed in square brackets in or-der not to lose that information.

When a lectotype has been designated, the type locality isthe locality of the lectotype. The original type locality [thesum of the localities of all the syntypes mentioned in theoriginal description], is not listed, unless justified.

Linnean species, pre-Linnean literature, unpublishedsources. — The identity and synonymy of species named


9

by Linnaeus [Linné] and other early authors present partic-ular problems, since many of them named species not onthe basis material they personally examined but by referenceto earlier literature sources. For example, in his Systemanaturae, Linnaeus (1758) based most species names on ear-lier accounts by himself and others. These have been tracedwhen possible and the exact bibliographic references given.These secondary references have been examined too; veryoften this actually creates additional problems because thesesecondary authors themselves refer to older publications, etc.I have not always included such secondary references andhave only rarely searched the tertiary and earlier sources.Additionally, several of these earlier works exist in variouseditions and Linnaeus' (and other's) bibliographic referencesare not detailed enough to decide which editions were used.For example, I have had the rare opportunity of examiningside by side different editions of Gesner's Nomenclator aqua-tilium and Fischbuch but could not find all texts to whichLinnaeus referred. As these books usually are considered tobe antiquities or collector-items, interlibrary loans or photo-copies are not possible. Examination and comparison of thevarious editions would mean travelling to a number of li-braries and investing a lot of time and money, beyond thelimits of the present work. Although there is an obvious his-torical and academic interest, the utility of the exercise is notobvious in the context of biological research and usable out-puts. In such instances, I merely list the reference as givenby the original author, updated into current bibliographic sys-tem. References to unpublished data are usually omitted un-less they are relevant for nomenclatural purposes [for exam-ple, reference to an unpublished figure of a type]. Type lo-calities listed are those given by the author of the new name;but the actual type locality is that listed by the author(s) onwhich the account is based. Where holotypes are extant, or iflectotypes or neotypes have been designated, the locality ofthese specimens of course becomes the type localities. Toidentify the type specimens of nominal species described bythese earlier authors, one has to follow about the same pro-cedure, that is, to find the specimens on which the accountscited by (e.g.) Linnaeus are based (for examples, see Wheel-er, 1958, 1985, 1991; Fernholm & Wheeler, 1983; Kottelat,2003c; Kottelat & Persat, 2005; Kottelat & Freyhof, 2009).Again, this is a tedious and time consuming task; I did searchsome such cases when this was essential for solving nomen-clatural problems, but did not search in detail all these cases.

(Contrary to a common belief, names created by Linnaeusare not sacred, however, since Systema naturae has beendesignated by later taxonomists as the starting point of to-day's nomenclatural system, the names he used are the firstavailable names for the concerned taxa. By the simple logicof the principle of priority they will remain, regardless ofhow usable or informative the descriptions are—and theyusually are useless without recourse to other sources).

Incertae sedis, genera inquirenda, species inquirendae,nomina dubia. — Incertae sedis are valid family, generaand species of uncertain taxonomic position. Genera incer-tae sedis are listed at the beginning of the Order or Familyto which they belong. For example, Pimelodus javus clear-

ly is a member of Siluriformes, but cannot be placed in anyfamily; it is therefore listed under Siluriformes, before thefamily accounts.

Species incertae sedis are listed at the beginning of the fam-ily to which they belong. Sometimes they are placed in thegenus in which my experience or that of knowledgeablecolleagues suggests they may belong. Alternatively, for spe-cies placed in genera to which there is a suspicion they donot belong, the generic name is placed in single quotationmarks (e.g. 'Genus' species), sometimes with a commentunder Taxonomic notes. The fate of a species incertae sedisis to be placed in a genus.

A species inquirenda (plural: species inquirendae) is a spe-cies of doubtful identity. Often they can be placed in a ge-nus but the description and the known material do not allowa decision as to whether or not the species is valid. Suchnames are listed immediately under the heading of the ge-nus to which they belong. Species inquirendae that cannotbe placed in any genus are listed immediately under the head-ing of the family to which they belong. A species inquiren-da may have great similarity to a valid species but its iden-tity may remain open to doubt; these are listed in the synon-ymy of that species, and are indicated by a question mark infront of the name. Some species inquirendae are poorly de-scribed but are nevertheless tentatively accepted as possi-bly valid, for example because an illustration in the originaldescription suggests they may be valid; awaiting confirma-tion or a usable description, they are listed as 'normal' spe-cies but with a question mark. It is noteworthy that a sub-stantial number of the taxa described from Vietnam in thelast 15 years falls into the category species inquirendae.

The fate of a species incertae sedis is that future studies willshow to which genus or family they belong. The fate of aspecies inquirenda is to be redescribed and either found tobe a valid species or a synonym of some other species.

A species inquirenda should not be confused with a nomendubium. A nomen dubium (plural: nomina dubia) is a nameof doubtful application that is impossible to link with aknown species, or that may apply to several species. Typi-cally, a nomen dubium would have been described in the18th or 19th century, with a few laconic sentences includ-ing no diagnostic characters usable today, or based on a paint-ing or on an artificially prepared specimen (examples in-clude species of Tetraodontidae based on deformed driedspecimens brought to Europe by seamen in the 18th centu-ry; or many species described from Chinese paintings in the19th century: these paintings usually were not based on agiven specimen but often were an artistic or idealised viewof the species, copied from earlier classical paintings, orsometimes simply imaginary). The fate of a nomen dubiumis not to remain so, but to become either a valid name or asynonym after either taxonomic examination or appropriatenomenclatural decisions.

A genus inquirendum (plural: genera inquirenda) is a ge-neric name that can be placed in a family but whose de-


10

scription and associated species (usually species inquirendaeor nomina dubia) do not allow a decision as to whether ornot it is valid. Such names are listed immediately under theheadings of the family to which they belong.

BASIC PRINCIPLES OF NOMENCLATURE

A sad reality is that a majority of the users of scientific names,especially those in the geographic area covered by this list,have not had the opportunity to study the rules of nomen-clature. This now also applies to most researchers complet-ing their studies in western countries. For this reason it seemsnecessary to start with a long introduction explaining basicprinciples of nomenclature, terminology, and how to under-stand the data in the present list. More experienced readerswill probably not need to read this section.

The most basic principle of nomenclature is that it dealswith only the names of organisms not with the organismsthemselves. The confusion between animals and their namesmars many taxonomic discussions and is becoming increas-ingly common and damaging. It is of concern that even theeditors of some scientific journals are no longer able to makethis distinction, especially in fashionable areas like molecu-lar systematics.

Nomenclature is about the correct formation and treatmentof names and the objective application of the 'legal' criteriaof a code, irrespective of taxonomic concepts or philosoph-ical approaches. Taxonomy is about the scientific study oforganisms and includes a level of subjective interpretationof observations that may differ among scientists.

Code. — Here, the word Code refers to the InternationalCode of Zoological Nomenclature. The current (4th) edi-tion was published in 1999 and superseded the previouseditions with effect from 1 January 2000. The Code is pub-lished under the responsibility of the International Com-mission of Zoological Nomenclature (ICZN, www.iczn.org),a body of zoologists (29 as of 2010), independent of politi-cal or national entities. Under exceptional circumstances andfollowing a prescribed procedure, the ICZN has the powerto suspend the application of any of the articles of the Code.These decisions (called Opinions and Directions) are pub-lished in the Bulletin of Zoological Nomenclature.

Nomenclatural acts. — A nomenclatural act is any of pub-lished act that affects the nomenclatural status of a scientif-ic name. This includes the creation of names, emendations,designation of types, rulings of the ICZN, etc. Nomencla-tural acts are valid if they satisfy the provisions of the Code;they are invalid and must be rejected if they do not followthe Code. Treating a name as a subjective synonym is a tax-onomic act, not a nomenclatural act.

Original descriptions. — Original description within themeaning of the Code (arts. 10–20) refers to the first use (cre-ation) of an available name.

Available name. — An available name is a name that satis-fies the criteria of the Code and may be used for a validspecies. The main criteria is that a new name must be ac-companied by a description and the designation of a name-bearing type (type specimen(s) for new species, type spe-cies for new genera; see below). An available name is notautomatically a valid name.

Since 2012, the Code allows the publication of new namesin electronic-only publications if they fulfill a number ofconditions. Among them, the work must have an ISSN orISBN number, be archived, and be registered in Zoobank(www.zoobank.org).

Valid name. — A valid name is the correct name applied toa species. To be valid, a name must first be available. But anavailable name is not automatically valid (junior synonymsare available names but invalid). A 'valid name' should notbe confused with a 'valid species'.

A nominal species is any of the available names created fora species, irrespective of its validity. If a valid species has xsynonyms, the valid name and the x synonyms are x+1 nom-inal species.

Species-group names, genus-group names, family-groupnames. — The species-group includes all the names of taxaof the rank of species and subspecies. The genus-group in-cludes all the names of taxa of the ranks genus and subge-nus. The family group includes all the names of taxa rankedabove the genus-group: superfamily, family, subfamily,tribes, etc.

Spellings. — A basic principle of nomenclature is that theoriginal spelling (the ones created by the author in the orig-inal description) must be retained. There are a few excep-tions and the Code is very precise about which spellingsmust be corrected (incorrect original spelling). There areno spelling that may be corrected, there are only spellingsthat must be corrected or that must not be corrected. Themain categories of corrections is that if the species name isa Latin adjective it must agree in grammatical gender withthe gender of the genus name. Incorrect original spellingsshould never be used. Corrections of incorrect original spell-ings allowed by the Code are called justified emendations.Any intentional (explained) corrections not allowed by theCode is an unjustified emendation and should never be used;unjustified emendations are available names with their ownauthor and date, are objective synonyms of the emendednames, may be homonyms and may be used as substitutenames. They are included in the list but I may have over-looked some. Any unexplained change or error is called anincorrect subsequent spelling. Incorrect subsequent spell-ings are not mentioned in synonymies, except if they haveused erroneously at least occasionally (example: Noema-cheilus and Nemachilus as commonly used incorrect spell-ings of Nemacheilus).

Date of publication. — While in everyday's language thepublication date is more or less equivalent to the date of


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printing, in the context of nomenclature, the Code definesthat the date of publication is the date at which a work couldfirst be obtained (for sale or by free distribution). The dateof publication of a work is important to determine prioritybetween two synonyms, two homonyms or two nomencla-tural acts. The Code rules that the earliest one has priorityover the youngest one (senior synonym vs. junior synonym;senior homonym vs. junior homonym). Even a difference ofone day is enough to give priority. The Code art. 23.9 al-lows exceptions (reversal of precedence), but only if veryprecise conditions are met.

The year of publication is retained as printed on the publi-cation. It may happen, however, that the work appeared at adate different from that printed on the publication. If a dif-ferent date is documented by reliable information, this datemust be retained for nomenclatural purposes. I have not at-tempted to check the effective publication date of all citedworks; this would have been very time and effort-consum-ing. I have invested time for such search only when it wasnecessary to establish the precedence between two works,or if it is was necessary to determine if a given work ap-peared before or after some of the dates important to theCode, or if there was a suspicion that a stated date is farfrom the actual date. Otherwise, I consider that usage of thedate printed on the publication is more important than theactual publication date (e.g. for retrieval on library shelvesor interlibrary loans).

Problems relating to dates associated with taxa are discussedunder Nomenclatural Notes. Those associated with specificpublications are mentioned in the Literature Cited section,under the respective titles. Some of the more complex casesare discussed separately: see Bibliographic Notes.

If nomenclatural acts are available from a valid electronic-only publication, the date of publication is the date of firstdistribution. New names and nomenclatural acts first madeavailable in electronic publications as 'accepted manuscript'or 'uncorrect proofs' are not available. Taxonomists shouldnot circulate manuscripts or proofs because this is a serioussource of future problems.

Priority, precedence. — There is a subtle difference be-tween priority and precedence. Priority indicates seniority,that a work, a name or a nomenclatural act was publishedbefore another one. Precedence indicates that a name mustbe used instead of another, either by application of the prin-ciple of priority, or because precedence is reversed for ex-ceptions prescribed by the Code or by rulings of the ICZN.

Authorship. — The names of animals are usually followedby the citation of their 'author'. For example the name of thecarp is Cyprinus carpio Linnaeus, 1758, in which Linnaeusis the author of the name C. carpio and 1758 the date of theoriginal description. The citation of the author and year ismerely a bibliographic reference. It is in no case an indica-tion of ownership. Unfortunately, many scientists forget orignore the purpose of the mention of the author's name andthis has sometimes resulted in unjustified emotional reac-

tions of authors when one of 'their' species is treated as aninvalid synonym by others. Somehow, naming a new spe-cies is the formulation of an hypothesis and there is no shameif an hypothesis is found to be erroneous. Vexed ego maynever accept a synonymy.

The name of an author is written in parentheses when thename is moved to another genus by subsequent authors. Forexample, Ompok bimaculatus (Bloch, 1794) was originallydescribed as Silurus bimaculatus by Bloch (1794).

For nomenclatural purposes, the Code (art. 50) defines theauthor(s) of a work as the person(s) who first published aname in a way that makes it available. In most cases theauthor of a name is the person whose name appears as theauthor of the book or article. In some cases of works bymore than one author, if one author only is responsible forthe name, then that person is author of the name and thename is then cited in the format Barbus binotatus Valenci-ennes, in Cuvier & Valenciennes, 1842. I personally thinkthat this kind of citation of authorship is contrary to the pur-pose of mentioning authors only as a bibliographic refer-ence but introduces a ownership aspect; in this example itseems even less desirable since the authors of the work hadthemselves decided that the work would appear under bothnames. In my eyes it serves no nomenclatural purpose tosearch who did what in such a work (of course I understandthe historical interest). But the Code says so and even theICZN has ruled so on this precise case.

If it is established that a person other than the person namedas the author(s) of a work is alone responsible for both thename and the conditions making the name available, thenthat person is the author of the name (often called second-ary author). For example, in Schneider (1801), the nameand the description of Mugil cirrhostomus are from unpub-lished notes of Forster. Schneider had no specimen and noother source to describe the species; therefore Forster isauthor of the name, and the name is cited as M. cirrhosto-mus Forster, in Schneider, 1801. [Forster's manuscript waslater published by Lichtenstein (1844) and the two texts canbe compared.]

If an external person is author of the description (for exam-ple personal notes) and the named author of the work citeshim and proposes a name for the taxon, then the named au-thor of the work remains author of the name. For example,Schneider (1801) used some of Forster's descriptions butchoose other names (for example to avoid homonymy); inthis case, Forster is not responsible for both the name andthe conditions making it available, and therefore Schneideralone is the author.

If an external person merely suggested a name for a speciesdescribed in a work, this does not make him the author ofthe name. The author of the work is responsible for the con-ditions making the name available (i.e., the description, des-ignation of types, etc.) and therefore is the sole author. Forexample, when Valenciennes (1846) described Cobitis fas-ciata he commented that in their notes the collectors (Kuhl


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and van Hasselt) had a drawing of this fish, which they hadlabelled Naemacheilus fasciatus. Kuhl and van Hasselt trag-ically died before they could publish the description. Twen-ty-four years later, Valenciennes used the name for the newspecies and wrote the description himself. This makes himalone the author. The species must be cited as C. fasciataValenciennes, in Cuvier & Valenciennes, 1846 and not C.fasciata Kuhl & van Hasselt, in Cuvier & Valenciennes, 1846.

Posthumous works belong to these categories but can bemore complex. Sometimes it is simply a matter of some-body publishing a completed manuscript by a deceasedfriend or colleague; in such case, the author of the manu-script is clearly the author of the work and of the names.The Code art. 50.1.1 explicitly mentions "satisfying the cri-teria of availability other than actual publication". This meansthat if both the names and the descriptions of the new taxaare entirely the work of a deceased person, then he is theauthor. To organise the actual publication is excluded fromthe conditions of authorship.

If the description of a given species has been modified bythe editor, both original writer and editor might be co-au-thor of the name. If the description is completely rewrittenby the editor, based on his own observations, then he is theauthor of the name.

Examples: Forsskål died during a travel around the Red Sea.His notes were later assembled, organised and published byNiebuhr, using the names in Forsskål's notes and withoutwork on the content of the text. Forsskål is author of thenames. For bibliographic (librarian) purposes, he is also treat-ed as author the work (see also Bibliographic Notes).

The manuscript of Bloch's Systema ichthyologiae was to-tally rewritten by Schneider, who also added numerous spe-cies, the index etc. Schneider is author of the work, as indi-cated on title page, but Bloch is author of some taxa.Schneider explicitly indicated the species he described.

When he died, Spix had not written the text on the fishes hecollected in Brazil. He had supervised the preparation ofmost plates and had named the species on the plates. Agas-siz was hired to write a text that could be distributed withthe plates. He wrote the descriptions of all species. For somehe ignored the names on the plates and created new names,and he is author of these names. For the species names thatappear only in plates, created by Spix, because the platesalone are among the conditions sufficient to make a name

available, Spix is the author. And in the cases Agassiz wrotedescriptions and used the names created by Spix on theplates, they are coauthors of the name.

Type species. — Each genus-group name has a type spe-cies. The type species of a genus name is the species whosename determines the validity of a genus. If several speciesare placed in genus X, with type species Xx and this genusis later divided into two genera, the genus which includespecies Xx will continue to be genus X while the other ge-nus will have another name. If two genera have the sametype species, they are objective synonyms. Genus-groupnames proposed after 1930 without the fixation of a typespecies are not available (Code art. 13.3) [note that art. 13.3requires that the fixation be "in the original publication [Art.68]" and that art. 68 includes, as "type species fixed in theoriginal publication", those established by original designa-tion, by monotypy, by absolute tautonymy and by Linneantautonymy].

Type genus. — Each family-group name has a type genus.The type genus is the genus whose name has been used toform the name of the family. For example Silurus is the nameof the genus used to form the family group names Siluridae,Siluriformes, Silurinae, etc. For nomenclature purposes thesethree words are a single name. Whatever the rank within thefamily-group, these names retain the same author and date.

Type specimens. — Each species-group name has a type.The type of a species name is the specimen on which thename is based; the phrase name-bearing type is more ap-propriate but, in order to simplify texts, is not usually used.The type specimen is the type of a name, not of a species. Itis therefore erroneous to understand the type as a 'model'representation of a species or a specimen to which all spec-imens must be identical to be called the same species. Thetype concept is a nomenclatural standard and totally inde-pendent of any taxonomic judgements or philosophical the-ory. The type is only used to objectively define to whichspecies the name must be applied. If the type specimen ofthe name Yus belongs to species 1, then the name of species1 is Yus. If the type specimens of the names Yus and Xusbelong to species 1, then the names Yus and Xus are syn-onyms (and the senior one has priority).

Only primary types (name bearing types) are listed here.Primary types are holotypes, lectotypes, neotypes and syn-types. Other type categories recognised by the Code areparatypes and paralectotypes but have no nomenclaturalfunction. Other 'type' categories (e.g. allotypes, topotypes,paratopotypes, paraneotypes) are not recognised by theCode, should not be used and are ignored here. Among them,allotype is sometime used to designate one of the paratypesof a sex different from that of the holotype; topotype is usedas a shortened way to say 'a specimen collected at the local-ity where the primary type was collected'.

The holotype is the specimen that has been explicitly desig-nated so (or by a similar wording) in the original descrip-tion by the original author, or the only specimen available

author author of and name then authorof work conditions itself of name is

is making name created byavailable is

A A A AA A B AA B A AA B B B, in AA A and B B A & B, in AA A and B A A & B, in A


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to the author, if there is clear evidence that the author basedthe nominal species on a single specimen. There is only oneholotype per species. In all cases where there is clear evi-dence that the author based the nominal species on morethan one specimen (including literature records) but did notdesignate a holotype, then all these specimens are syntypes.When it is not possible to determine from the original de-scription if a name is based on one or several specimens,I usually use 'types' or 'holotype?'.

If the nominal species is based on a specimen explicitly des-ignated as holotype and a number of additional specimensare also explicitly designated as types, these are paratypes;allotypes are thus paratypes. The sum of the holotype+para-types or the sum of the syntypes is called the type series. Insome cases (especially for species described by P. Bleeker),I indicate the size ranges of the type series in square brack-ets since this can be an important tool to recognise them(example: syntypes [12, 41–43 mm SL]).

In cases where there is no holotype but only a series of syn-types, one of the syntypes may be designated as lectotype; itthen has the same value as the holotype. The remaining syn-types then become paralectotypes and lose their status asprimary types. Lectotypes are designated when it is demon-strated or suspected that the type series includes more thanone species; it allows the name to be definitively fixed tothe nominal species to which the lectotype belongs. Inci-dentally, the designation of a lectotype also restricts the typelocality to the locality of the lectotype, excluding the local-ities of the other syntypes. Paratypes and paralectotypes arenot listed hereunder as they are not name-bearing types.

If none of the specimens of the original type series remains,or if the holotype or lectotype no longer exist (they have notbeen preserved, are lost, or destroyed) and if the name can-not be unambiguously linked to a valid species, then (andonly then) a specimen can be designated as neotype thatwill have the same function as the holotype. Incidentally,the designation of a neotype also restricts the type localityto the locality of the neotype. All designations of neotypesthat do not fully satisfy these and several other conditionslaid down in the Code are invalid and must be ignored.

A number of neotype designations are invalid because theneed for a neotype is not stated or demonstrated. This re-quirement did not exist in the 1985 Code (art. 75(b)) and animplicit justification was enough. The requirement becameexplicit in the 1999 Code, with an added clause (art. 75.3.1)requiring "a statement that [the neotype] is designated withthe express purpose of clarifying the taxonomic status orthe type locality of a nominal taxon". Unfortunately the 1999Code is missing a clause explaining what should be done ofneotype designation validly made before 2000 but invalidunder the 1999 Code because of the absence of the state-ment. Formally, they become invalid because art. 86.3 statesthat all former editions of the Code have no force.

This I interpret as an oversight of the editors of the 1999Code and is unintentional, otherwise it could be the cause

of very serious instability in some groups, which would betotally against the spirit of the Code. Many of the pre-2000neotype designations do not have the statement required bythe 1999 Code. I have retained as valid the neotypes validlydesignated before 2000 under the 1985 Code.

After the original description, it may be necessary for laterauthors to re-examine the primary type of a nominal speciesin order to decide to which taxonomic species it applies, forexample in cases when several similar species are later dis-covered and the original description does not mention thecharacters now decisive to determine to which of these spe-cies the name must be applied. It is, however, not a necessi-ty to examine a primary type if the original description pro-vides all the information needed for identification. In fact,types may be fragile specimens, and they should not be han-dled if not justified, and persons without experience shouldnot be permitted to handle them. Primary types must bedeposited in museums or other responsible institutions andwith staff able to conserve them and make them accessibleto later researchers. Even if there is political pressure in somecountries to consider types as national property, types donot belong to a country but to science and must be accessi-ble to competent scientists irrespective of their nationality.Neotypes, by definition, must be deposited in a recognisedinstitution (e.g. museum).

A number of species described by earlier authors do not haveknown types or they have been lost since the original de-scription. This does not affect the availability of a name.For example, a specimen described in the field and latereaten by an author remains the type specimen. Or a speci-men used as model for a figure remains the type specimen,even if it has not been preserved.

When known, institutions in which primary types are de-posited are listed, together with register number and, whenknown, the number of specimens in square brackets (exam-ple: AAA 1234 [2], BBB 1233 [1]). When the primary typeswere deposited in a collection but cannot presently be locat-ed, the institution is listed as they may still be present (mis-identified, misplaced, uncatalogued), or as a starting pointfor further search. The source for the catalogue number isgiven when it is not the original description; besides, manyof those listed in the original descriptions have also beenchecked in published catalogues or in the institutions them-selves. When there is a series of syntypes, I listed thoseI could trace in the literature, but made no effort to trace thewhole series; this would have been tedious, many of themhaving possibly been used for exchanges between institu-tions, etc. NT indicates that there is no (or apparently no)preserved type material, LU that there was apparently pre-served type material but that its whereabouts are not known.A question mark in front of the abbreviation of an institu-tion indicates that the type(s) is possibly there or that thetype status of the specimen is not certain.

Institutional abbreviations used in the text are listed below.For institutions for which no abbreviations have been usedin literature, the abbreviations follow current use by work-


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ers at these institutions (where possible) or Leviton et al.(1985; Leviton & Gibbs, 1988) or Eschmeyer (2010). I didnot automatically follow Leviton et al. and Eschmeyer as astandard because for non-US collections the abbreviationsthey list are often not those actually used by the institutionsthemselves. In case the abbreviations used in these lists dif-fer from those used by workers at these institutions, I retainthe second one (as long as they make sense and do not rep-resent personal, bureaucratic or chauvinistic idiosyncrasy).

Synonyms. — The word synonym is used with the meaningit has in the Code, that is a new name applied to a speciesthat already had a name. Names erroneously used for a spe-cies other than the one originally described under that nameare misidentifications. Misidentifications are not synonymsand are not included in this catalogue. The Code rules thatin case an author thinks that two names actually refer to asingle species (i.e., they are synonyms), the name publishedfirst (senior synonym) is the valid name; the name publishedlater (junior synonym) is invalid (cannot be used). The jun-ior synonym nevertheless remain available; should a laterauthor find that the type specimens of the two names actual-ly refer to different species, the junior synonym might beused again (if it satisfies conditions set by the Code).

If the two synonyms are based on the same specimen (i.e.,they have the same primary type), they are objective syn-onyms and the junior synonym is invalid. If the two namesare based on different primary types that an author consid-ers as belonging to a single species, they are subjective syn-onyms (they are subjective because this is the taxonomicjudgement of an author and other authors may disagree; inthe opposite case, objective synonymy is a purely nomen-clatural issue, not depending on of taxonomic judgement).

Homonyms. — Two available names with identical spell-ings and created independently for different taxa are calledhomonyms. The Code rules that the name published first(senior homonym) is the valid name; the name publishedlater (junior homonym) cannot be used and must be replaced.Junior homonyms are permanently invalid, unless satisfy-ing some precise conditions of the Code.

In the species-group, two homonyms created in the samegenus are called primary homonym. Example: Barbus yun-nanensis Fowler, 1958 is a junior primary homonym of Bar-bus yunnanensis Regan, 1904.

Two species names originally established in different gen-era but later combined with the same genus name are calledsecondary homonyms. Example: Crayracion fluviatilis var.ocellata Steindachner, 1870 was not a homonym of Tetra-odon ocellatus Linnaeus, 1758 when established. Later (in1975), it was treated as a valid species of Tetraodon and itsname became T. ocellatus (Steindachner, 1870), a juniorsecondary homonym of T. ocellatus Linnaeus, 1758.

Replacement names. — If a species name becomes invalidbecause it is a junior secondary homonym, it must be re-placed. The name used for replacement is called substitute

name. The substitute name is the next oldest synonym. Ifthere is no available synonym, then a new replacement nameis established. The new replacement name takes the sametype as the replaced name. In the example above, Dekkers(1975) treated Crayracion ocellatus as a valid species ofTetraodon and made it a junior homonym of Tetraodon ocel-latus. He created the new replacement name T. steindachnerito replace the junior homonym.

A junior secondary homonym rejected and replaced before1961 is definitively invalid (there may be exceptions; Codeart. 59.3). But a junior secondary homonym rejected after1960 but later considered to be in a genus different from thesenior homonym becomes valid again (Code art. 59.4). Inthe above example,when Crayracion ocellatus Steindachner,1870 was treated as a valid species of Tetraodon it had to bereplaced by Tetraodon steindachneri Dekker, 1975. ButT. steindachneri is now considered to be a valid species ofDichotomyctere and the senior synonym must be reinstatedand the valid name is now D. ocellatus (Steindachner, 1870),not D. steindachneri.

Occasionally, some authors have replaced names becausethey overlooked an already available name that should havebeen used as substitute name, or because they did not likean existing name, or because they found it inappropriate, orto follow the nomenclature rules at the time, or under politi-cal pressure. These replacement names are invalid and can-not be used. These names are called unnecessary replace-ment names. In the above example, after the creation of thenew replacement name Tetraodon steindachneri Dekkers,1975 it was discovered that the misidentified T. biocellatusTirant, 1885 in fact was also a junior synonym of T. ocellatus(Steindachner, 1870). Therefore T. biocellatus became thevalid substitute name for T. ocellatus (Steindachner, 1870)and T. steindachneri became a junior synonym ofT. biocellatus. An later, after moving the species to Dycho-tomyctere as mentioned above, the replaced secondary jun-ior homonym T. ocellatus was reinstated as D. ocellatus(Steindachner, 1870).

Infrasubspecific names. — Infrasubspecific names arenames originally intended for categories below the subspe-cies level (for example: varieties, natios). These names arenot recognised as valid by the Code. They are neverthelesslisted here. For infrasubspecific names, I have indicated thelocality stated by the original author, but have not listedmaterial. Even if these have sometime been called type lo-calities and types in the literature, as the names are not avail-able for zoological nomenclature, these 'type localities' and'types' have no nomenclatural status.

Infrasubspecific names may become validated by a subse-quent use as subspecies or species name. Whenever I foundan infrasubspecific name validated this way I noted it; ifsuch subsequen

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