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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



There are 3108 valid and named native fish species in the inland waters of Southeast Asia between the Irrawaddy and Red River drainages, the small coastal drainages between the Red River and Hainan, the whole Indochinese Peninsula, Andaman and Nicobar Islands, Indonesia (excluding Papua Province, Waigeo, Aru [but Kai is included]), and the Philippines. They belong to 137 families. Their taxonomy and nomenclature are reviewed. The original descriptions of all 7047 recorded species-group names and 1980 genus-group names have been checked in the original works for correct spelling, types, type locality and bibliographic references. The bibliography includes about 4700 titles. Synonymies are given, based on published information as well as unpublished observations. The names of 49 introduced species and 347 extralimital taxa cited in the discussions have also been checked. The original descriptions of all species not present in the covered area but cited as type species of genera have been checked for availability, authorship, date and correct spelling. The availability of some family-group names has been checked when there was suspicion of possible nomenclatural problems. Bibliographic notes include new informations on the dates of publication of works by, among others, Bleeker, Bloch, Heckel and Steindachner and discussion of authorship of names in various works.
An International Journal of Southeast Asian Zoology
Supplement No. 27 22 November 2013
The Raffles Bulletin
of 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,
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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
(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 ies (Diptera:
Tephritidae) in Southeast Asia. Raf es Bulletin of Zoology, Supplement 7: 1–92.
8. Ng, P. K. L. & K. S. Tan (eds.), 2000. The biodiversity of South China Sea. Raf es Bulletin of Zoology, Supplement 8: 1–673.
9. Kurahashi, H. & F. R. Magpayo, 2000. Blow ies (Insecta: Diptera: Calliphoridae) of the Philippines. Raf 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. Raf es
Bulletin of Zoology, Supplement 10: 1–187.
11. Ng, P. K. L., D. Wowor & D. C. J. Yeo (eds.), 2002. Scienti c results of the Anambas Expedition 2002. Raf 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.
Raf es Bulletin of Zoology, Supplement 12: 1–434.
13. Kottelat, M. & D. C. J. Yeo (eds.), 2005. Southeast Asian freshwater sh diversity. Raf es Bulletin of Zoology, Supplement 13:
14. Tan, H. & R.-Q. Jan (eds.), 2007. Proceedings of the 7th Indo-Paci c Fish Conference. Raf es Bulletin of Zoology, Supplement
14: 1–434.
15. Wang, L. K. & C. J. Hails, 2007. An annotated checklist of birds of Singapore. Raf es Bulletin of Zoology, Supplement 15:
16. Tan, S. H. & P. K. L. Ng (eds.), 2007. Crustacean Supplement 1. Raf 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. Raf 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. Raf
es Bulletin of Zoology, Supplement 18: 1–264.
19. Tan, S. H. & I.-S. Chen (eds.), 2008. Aquatic biodiversity of the South China Sea. Raf es Bulletin of Zoology, Supplement 19:
20. Tan, S. H. & M. E. Y. Low (eds.), 2009. Crustacean Supplement II. Raf 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 classi cation of living and fossil genera of decapod crustaceans. Raf es Bulletin of Zoology, Supplement
21: 1–109.
22. Tan, K. S. (ed.), 2009. Fourteenth International Marine Biology Workshop 2006: The marine ora and fauna of Singapore.
Raf 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. Raf es Bulletin of Zoology,
Supplement 23: 1–181.
24. Davison, G. W. H. & C. S. W. Chia (eds.), 2011. Proceedings of the fth International Hornbill Conference, Singapore, 22–25
March 2009. Raf 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). Raf 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).
Raf es Bulletin of Zoology, Supplement 26: 1–199.
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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. Raf es Bulletin of Zoology, Supplement
1: 1–128.
2. Deeleman-Reinhold, C. L., 1995. The Ochyroceratidae of the Indo-Paci c region (Araneae). Raf 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. Raf 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. Raf es Bulletin of Zoology, Supplement 4: 1–141.
5. Kurahashi, H., N. Benjaphong & B. Omar, 1997. Blow ies (Insecta: Diptera: Calliphoridae) of Malaysia and Singapore. Raf 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. Raf es Bulletin
of Zoology, Supplement 6: 1–288.
(continues on back cover)
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The Raf es Bulletin of Zoology Raf es Museum of Biodiversity Research Department of Biological Sciences National
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ISBN 978-2-8399-1344-7
THE RAFFLES BULLETIN OF ZOOLOGY 2013 Supplement No. 27: 1–663
Date of Publication: 22 Nov.2013
© National University of Singapore and the author
Maurice Kottelat
Case 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
Abstract ......................................................................... 3
Introduction ................................................................... 4
Basic principles of nomenclature................................ 10
Abbreviations used...................................................... 15
Class Chondrichthyes
Subclass Elasmobranchii
Division Selachii
Order Orectolobiformes
Family Hemiscylliidae .......................... 18
Family Stegostomatidae........................ 19
Order Carcharhiniformes
Family Carcharhinidae ......................... 20
Division Batoidea
Order Pristiformes
Family Pristidae .................................... 23
Order Rajiformes
Family Rhinobatidae............................. 24
Order Myliobatiformes
Family Dasyatidae ................................ 25
Family Potamotrygonidae..................... 28
Family Myliobatididae.......................... 29
Class Actinopterygii
Division Teleostei
Order Osteoglossiformes
Family Osteoglossidae ............................... 30
Family Arapaimidae ................................... 31
Family Notopteridae ................................... 31
Order Elopiformes
Family Elopidae ......................................... 33
Family Megalopidae ................................... 33
Order Albuliformes
Family Albulidae ........................................ 34
Order Anguilliformes
Family Anguillidae ..................................... 37
Family Moringuidae ................................... 39
Family Muraenidae .................................... 41
Family Ophichthidae .................................. 43
Family Muraenesocidae ............................. 48
Order Clupeiformes
Family Pristigastridae ................................. 49
Family Engraulididae ................................. 52
Family Chirocentridae ................................ 57
Family Clupeidae ....................................... 58
Family Sundasalangidae............................. 64
Order Gonorhynchiformes
Family Chanidae......................................... 64
Order Cypriniformes
Family Cyprinidae ...................................... 65
Family Psilorhynchidae ............................ 171
Family Gyrinocheilidae ............................ 173
Family Botiidae ........................................ 173
Family Vaillantellidae............................... 176
Family Cobitidae ...................................... 176
Family Ellopostomatidae.......................... 184
Family Barbuccidae.................................. 185
Family Balitoridae .................................... 185
Family Gastromyzontidae ........................ 191
Family Serpenticobitidae ......................... 197
Family Nemacheilidae.............................. 198
Order Characiformes
Family Characidae.................................... 215
Family Serrasalmidae ............................... 215
Order Siluriformes
Family Loricariidae .................................. 215
Family Amblycipitidae ............................. 216
Kottelat: Inland fishes of Southeast Asia
Family Akysidae ....................................... 217
Family Sisoridae ....................................... 221
Family Cranoglanididae ........................... 232
Family Siluridae ....................................... 232
Family Chacidae ....................................... 240
Family Plotosidae ..................................... 240
Family Clariidae ....................................... 242
Family Ariidae .......................................... 245
Family Schilbeidae ................................... 251
Family Pangasiidae .................................. 253
Family Bagridae ....................................... 256
Order Osmeriformes
Family Plecoglossidae .............................. 268
Family Salangidae .................................... 268
Order Salmoniformes
Family Salmonidae ................................... 270
Order Aulopiformes
Family Synodontidae................................ 270
Order Gadiformes
Family Bregmacerotidae .......................... 271
Order Ophidiiformes
Family Carapidae ..................................... 271
Family Bythitidae ..................................... 272
Order Batrachoidiformes
Family Batrachoididae ............................. 272
Order Lophiiformes
Family Antennariidae ............................... 274
Order Mugiliformes
Family Mugilidae ..................................... 275
Order Atheriniformes
Family Telmatherinidae ............................ 283
Family Phallostethidae ............................. 284
Family Atherinidae ................................... 286
Order Beloniformes
Family Adrianichthyidae .......................... 288
Family Hemiramphidae ............................ 290
Family Zenarchopteridae ......................... 293
Family Belonidae ..................................... 297
Order Cyprinodontiformes
Family Aplocheilidae ............................... 299
Family Poeciliidae .................................... 300
Order Beryciformes
Family Holocentridae ............................... 301
Order Gasterosteiformes
Family Indostomidae ................................ 301
Family Syngnathidae ................................ 301
Order Synbranchiformes
Family Synbranchidae .............................. 307
Family Chaudhuriidae .............................. 309
Family Mastacembelidae.......................... 310
Order Scorpaeniformes
Family Scorpaenidae ................................ 314
Family Tetrarogidae ................................. 315
Family Synanceidae ................................. 316
Family Platycephalidae ............................ 316
Order Perciformes
Suborder Percoidei
Family Ambassidae ............................. 318
Family Latidae .................................... 323
Family Lateolabracidae ...................... 324
Family Percichthyidae ........................ 324
Family Serranidae ............................... 325
Family Pseudochromidae ................... 328
Family Opistognathidae...................... 329
Family Centrarchidae.......................... 329
Family Apogonidae............................. 329
Family Sillaginidae ............................. 330
Family Carangidae .............................. 331
Family Leiognathidae ......................... 337
Family Lutjanidae ............................... 340
Family Datnioididae ........................... 344
Family Lobotidae ................................ 344
Family Gerreidae ................................ 345
Family Haemulidae ............................. 347
Family Nemipteridae .......................... 349
Family Lethrinidae.............................. 350
Family Sparidae .................................. 352
Family Polynemidae ........................... 353
Family Sciaenidae ............................... 356
Family Mullidae .................................. 360
Family Monodactylidae ...................... 362
Family Toxotidae ................................ 362
Family Drepaneidae ............................ 365
Family Chaetodontidae ....................... 365
Family Nandidae ................................. 366
Family Pristolepididae ........................ 366
Family Badidae ................................... 367
Family Terapontidae ........................... 368
Family Kuhliidae ................................ 371
Suborder Labroidei
Family Cichlidae ................................. 373
Family Pomacentridae ........................ 375
Family Labridae .................................. 379
Family Scaridae .................................. 380
Suborder Trachinoidei
Family Trichonotidae.......................... 382
Suborder Blennioidei
Family Blenniidae ............................... 382
Suborder Callionymoidei
Family Callionymidae ........................ 385
Suborder Gobioidei
Family Rhyacichthyidae ..................... 386
Family Odontobutidae ........................ 387
Family Eleotrididae ............................ 388
Family Kraemeriidae .......................... 396
Family Gobiidae ................................. 396
Family Amblyopidae .......................... 432
Family Ptereleotrididae....................... 435
Suborder Kurtoidei
Family Kurtidae .................................. 436
Suborder Acanthuroidei
Family Ephippidae .............................. 437
Family Scatophagidae......................... 438
Family Siganidae ................................ 439
Family Acanthuridae........................... 441
Suborder Scombroidei
Family Sphyraenidae .......................... 444
Family Scombridae ............................. 446
Suborder Anabantoidei
Family Anabantidae ............................ 446
Family Helostomatidae ....................... 447
Family Osphronemidae....................... 448
Suborder Channoidei
Family Channidae ............................... 458
Order Pleuronectiformes
Family Paralichthyidae............................. 462
Family Tephrinectidae .............................. 463
Family Soleidae ........................................ 463
Family Cynoglossidae .............................. 466
Order Tetraodontiformes
Family Triacanthidae................................ 468
ABSTRACT. — There are 3108 valid and named native fish species in the inland waters of
Southeast Asia between the Irrawaddy and Red River drainages, the small coastal drainages
between the Red River and Hainan, the whole Indochinese Peninsula, Andaman and Nicobar
Islands, Indonesia (excluding Papua Province, Waigeo, Aru [but Kai is included]), and the
Philippines. They belong to 137 families. Their taxonomy and nomenclature are reviewed. The
original descriptions of all 7047 recorded species-group names and 1980 genus-group names
have been checked in the original works for correct spelling, types, type locality and bibliographic
references. The bibliography includes about 4700 titles. Synonymies are given, based on
published information as well as unpublished observations.
The names of 49 introduced species and 347 extralimital taxa cited in the discussions have
also been checked. The original descriptions of all species not present in the covered area but
cited as type species of genera have been checked for availability, authorship, date and correct
spelling. The availability of some family-group names has been checked when there was suspicion
of possible nomenclatural problems.
Bibliographic notes include new informations on the dates of publication of works by, among
others, Bleeker, Bloch, Heckel and Steindachner and discussion of authorship of names in various
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,
– 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, Cyprinus
lamta, Engraulis rhinorhynchos, Esox alepidotus, Esox argenteus Gmelin, Esox argenteus
Schneider, Equula longispinis, Gobiomoroides piso, Gobius niger, Gonorhynchus bimaculatus,
Hemiramphus buffonis, Hemiramphus brevirostris, Hemiramphus georgii, Hemiramphus
russelli Valenciennes, Johnius cataleus, Lobotes auctorum, Neostethus borneensis,
Parosphromenus parvulus Foersch & Korthaus, Pellona leschenaulti, Raja edentula, Raja
guttata Shaw, Raja narinari, Rasbora trilineata, Synaptura achira, Teuthis brevirostris, Teuthis
– neotype designation for: Engraulis dussumieri, Lutjanus gymnocephalus, Ovoides fasciatus,
Ovum commersoni, Platygaster megalopterus, Scarus schlosseri, Sciaena jaculatrix, Sciaena
– declaration as nomina protecta: Albula Scopoli, Aplocheilus, Clupea quadrimaculata, Cyprinus
bola, 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, Raja
ommescherit, Raja scherit, Raja schoukie, Sphyraena japonica Bloch, Toxotes microlepis
Family Monacanthidae ............................. 469
Family Tetraodontidae.............................. 469
Family Molidae ........................................ 481
Appendices ................................................................ 481
Acknowledgements ................................................... 484
Bibliographic notes.............................................. 485
Literature cited..................................................... 505
Kottelat: Inland fishes of Southeast Asia
– 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, Apistus
longispinis over A. bougainvillii, Barbus balleroides over B. hypsylonotus, Carassioides
macropterus 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, Placocheilus
bibarbatus 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 been
generally ignored.
– Oxygastri of Bleeker (1860c) is not available because it is a descriptive term, and not based on
the genus name Oxygaster;
– new genera: Desmopuntius (type species: Barbus hexazona Weber & de Beaufort, 1912),
Oliotius (type species: Capoeta oligolepis Bleeker, 1853), Puntigrus (type species: Barbus
partipentazona 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
Nowadays it seems fashionable among many zoologists, botanists, anatomists
and 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 opposite
error. 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 of
the doubt. The now fashionable discussions about cryptic
species do not change the situation: one nucleotide does not
make a species, be it ever so cryptic. Further, in fishes, the
cryptic species discovered by molecular techniques that I
have 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' fish
species that have been described from Vietnam in recent
years. Their description is of a quality that makes it simply
impossible 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 times
are not recognised. Cladistic molecular phylogeny (which
uses principles and mathematical algorithms that were called
phenetics 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 two
papers (Mayden & Chen, 2010; Tang et al., 2010; see Britz
& Conway, 2011a–b). This shows that it is imprudent to
instantly adopt the latest theory and that naming every little
temporary lineage uncovered by molecular analysis has lit-
tle justification.
For this catalogue I have examined personally the original
descriptions of all the species and genera recorded in the
inland waters of Southeast Asia, all their synonyms (mak-
ing a total of about 7047 nominal species and about 1980
nominal genera). I also examined the original descriptions
of the type species of all genera if they were not known in
the area, and those of the 347 taxa cited in the Taxonomic
and Nomenclatural Notes but not present in area. All cited
nomenclatural acts were checked. The availability of all non-
fish names cited as senior synonyms of fish genera was
checked. Synonyms based on fossil taxa and which have
never been used for recent taxa are not included.
The present catalogue aims to present the state of the art of
our knowledge of the diversity of freshwater fishes of South-
east Asia. Work began in 1986 when I compiled a list of the
freshwater fishes of the Indochinese region (Kottelat, 1989).
The list expanded when I worked on a book on the fishes of
western 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 book
required the inclusion of all species that had been recorded
in inland waters, that is, including estuaries, most mangroves,
etc. In September 2013, the list includes 3107 valid native
species, in 707 valid genera and 137 families. Only named
species are included. I am aware of about 300 species to be
named soon or already on museum shelves and a fair num-
ber of synonyms to re-validate. I expect an additional 500
species still awaiting discovery in the wild. In addition, there
are 49 introduced and established species.
This catalogue is not the ultimate inventory of the fishes of
Southeast Asian inland waters. Many discoveries are still
ahead of us and a great amount of work remains to be done
before 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 of
living organisms and their phylogenetic relationships. The
other 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 and
their identification now, not sometime in the distant future.
If definitive conclusions are not possible with the available
data, 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, the
precautionary approach would be to retain them as distinct
awaiting (possible) further research.
This precautionary approach, however, has its limits. The
development of molecular techniques has led some to rec-
ognise as 'species' populations distinguished only by a few
nucleotides; complex statistics have been used to justify the
recognition of 'species' otherwise not distinguishable by
Kottelat: Inland fishes of Southeast Asia
About 6000 publications (in 22 languages) have been ex-
amined and about 4700 relevant ones are listed. All cited
references 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 of
an antique book) are marked in bold face (3 out of 4700
titles). A few references have been checked by trusted col-
After completion of the catalogue, all names have been cross-
checked against other databases. Especially, all names were
cross-checked against Eschmeyer's (2013) Catalog of Fish-
es (CoF) in 2001. For each entry in CoF disagreeing with
my 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 the
original literature was checked for a third time. Out of the
about 9785 checked names, the data (spelling, author, date,
types, type locality, etc.) for about 3440 differs from those
in CoF (35 %). Most of the differences are minor and of
little or no nomenclatural consequences (mainly related with
type localities), but a significant number of differences are
serious. The problems are more frequent with the pre-1860
non-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 made
about the quality of the data and there seem to have been
few or no efforts to carefully evaluate the contents of such
large databases. An analysis of the types of differences and
errors is in preparation.
A significant number of nomenclatural problems were dis-
covered and the application of the International Code of
Zoological Nomenclature (hereunder Code) results in a num-
ber of nomenclatural changes. In a few cases of changes
affecting family-group names or widely used names, requests
have been presented to the International Commission on
Zoological Nomenclature (ICZN) to retain these names in
their current usage (Scatophagidae, Ephippidae, Kottelat,
2010b; Siganidae, Kottelat, 2013b; Mystus, Kottelat & Ng,
2007). In other cases, I considered that the name changes
are minor and I simply applied the Code. At the genus level,
I consider that changes resulting from the application of the
Code do not create more problems than do changes result-
ing from the normal increase of our taxonomic knowledge
by the discovery of new taxa, new characters, etc. I consid-
er it appropriate to ask the ICZN to retain the current usage
in cases of potential confusion resulting from the discovery
of overlooked type species designations. But I consider it
unjustified 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 in
favour of reputedly 'well-known' names does not hold in a
geographic area where new discoveries still abound and
where the taxonomic system is still very unstable. Under
that logic, dozens of names should be suppressed and this
would affect the stability of nomenclature by making the
purpose of the Code irrelevant. Also, writing applications
for all these minor cases would mean more applications than
the ICZN receives in a year.
Limitations. — One of the limitations of this catalogue is
that I started it in 1986; the work spanned 24 years and four
operating 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, two
different editions of the International Code of Zoological
Nomenclature (ICZN, 1985, 1999) have been in use, which
differ slightly. I have tried to update all entries affected by
the changes but I may have missed some.
Another limitation is that the core target of this work and
my own experience is the 'real' freshwater fishes. My treat-
ment of these taxa is probably close to complete. But I am
likely to have missed some records of estuarine species, or
some 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 came
to the same situation with estuarine taxa, I tried to solve the
routine problems but decided not to address the more com-
plex ones. However, I discuss these cases and their possible
solutions where pertinent. The number of marine taxa with
nomenclatural problems was unexpectedly high, and many
well and long known genus and family names are involved.
A potential for small errors arose late in the preparation of
this catalogue. I had long tried to confirm or revise the chro-
nology of the many papers published by Pieter Bleeker and
it 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 in
different papers and journals. A consequence of this revised
chronology is that the now-established dates of availability
of many names differ from those commonly recognised, and
this has changed the precedence of the different description
of a few species, which now may have a different type se-
ries, or of some new genera, which now may have different
originally included species, thus potentially invalidating
earlier type species designations. I have tried to eliminate
this risk but I expect that some details would have escaped me.
Geographic and habitat coverages. — The geographic
coverage includes all inland water bodies of Southeast Asia
between (and including) the Kaladan, Irrawaddy and the Red
River drainages, the small coastal drainages between the Red
River and Hainan (included), the whole Indochinese Penin-
sula, Andaman and Nicobar Islands, Indonesia (excluding
Papua Province, Waigeo and Aru [but Kai is included]), and
the Philippines (Fig. 1).
All freshwater species are included. Introduced species that
became established are listed (marked by asterisks, *), but
without 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 the
records or identifications need critical reevaluations (which
was beyond the goals of this work). I preferred to be too
inclusive than too exclusive. For these species too, synony-
mies are complete and include the nominal species described
from freshwater as well as the marine ones; similarly, the
generic synonymies applying to these genera are complete.
Through a lack of familiarity with some groups I may have
overlooked some synonyms.
Listed names. — All known names are listed, including
infrasubspecific names, which are given in their original
form. The only names that are not included are those sub-
specific or infrasubspecific epithets typicus when they are
merely intended to denote the nominotypical subspecies;
such names are usually not nomenclaturally available and
should not be used.
Spellings. — The headings of all generic and specific ac-
counts have the correct spelling of all valid names. In the
synonymies, however, all names are given with their origi-
nal combination (except that interpolated subgeneric names
are 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 by
the 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 in
Nelson (2006), except within Cypriniformes, for which
I follow Šlechtová et al. (2007) and my personal experience.
When there is disagreement between authors with regard to
the 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 synonymies
of family-group names. Note that a family-group name keeps
its 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 or
family Leuciscidae.
Genera and species. — Genera are listed in alphabetical
sequence within family. Species are listed in alphabetical
sequence within genera.
Entries for genera include the valid name of the genus (in
bold, as a heading), the name of the genus with the spelling
in 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.
Kottelat: Inland fishes of Southeast Asia
is followed by information on possible subgeneric status in
the original description, type species, mode of designation,
information on possible nomenclatural acts associated with
the 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 (in
bold, as the heading), the name of the species as spelt in the
original description, the author, the year of publication, the
number 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 with
information on type locality and primary types, and infor-
mation on possible nomenclatural acts associated with the
name. If the name is based largely or totally on references to
the older literature, this information is listed first in the
renthesed block). This information is provided for all names
considered to be synonyms, in chronological sequence.
Additional information, if needed, is listed under Nomen-
clatural notes and Taxonomic notes. When names are cited
under Notes, which are not mentioned elsewhere in the text,
I usually (but not always) added the same data for that name
at the end of the paragraph, in brackets.
Transliteration of non-Latin alphabets. — Author names,
place names, and journal names in non-Latin alphabets, and
in 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 an
abstract, in a table of contents), the same spelling is used
here. 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 and
no 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 actually
complicate bibliographic search, especially for those not
familiar with these languages.
Unfortunately, some accents and diacritic marks may have
disappeared as standard western European keyboards and
software do not support them. This especially applies in the
case of the Vietnamese alphabet.
Type localities. — The type locality is the locality at which
the holotype, lectotype or neotype was collected. Although
mentioned in the Code, the type locality has no nomencla-
tural role. Simply, it is a convenient wording, it is shorter to
say type locality than 'the locality at which the primary name-
bearing type was collected', or to give the locality data in
In case there is no primary type but a series of syntypes
from different localities, the type locality is the sum of all
the localities of the syntypes, and all their localities are list-
ed (separated by a slash [ / ] where clarity requires it). In
very few cases (when the list of localities of syntypes is
very extensive), I have merely given a general description
of the localities. Localities are usually given with the origi-
nal spelling; this sometimes results in different spellings
being used for the same locality under different headings;
I have tried to introduce some consistency, but only in cases
where I was certain that the different spellings were really
referring to the same place, or when the different spellings
were used for the locality of the very same specimen, or
referring to the very same bibliographic source. Alternative
spellings or modern equivalents are given in square brack-
ets, but this has not been systematically attempted. Locality
descriptions 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 and
they have not been deleted in order to avoid ambiguities
when using local maps [but the word river, lake, etc. has
been added]. Local names have been used, except for a few
well known rivers and lakes with a common English name
used 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 political
entities (country, province, state, etc.) and river basin as an
aid to the reader. For larger topographic features that have
several different names, a single one has been consistently
used; this especially applies to those traversing different
countries (e.g. Mekong River); I usually retained the name
easiest to find for readers not familiar with local toponymy
or, when it exists, the English name used in international
literature (e.g. Salween River and not Nu Jiang, Salawin,
Thanlwin, Salouen, or fGyl mo rNGul chu [a transcription
from 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 slightly
different 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 for
infrasubspecific names based on aberrant specimens; they
are given only if the name has been created for a particular
geographical form.
When a neotype has been designated, the type locality is the
locality 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 is
the locality of the lectotype. The original type locality [the
sum of the localities of all the syntypes mentioned in the
original description], is not listed, unless justified.
Linnean species, pre-Linnean literature, unpublished
sources. — The identity and synonymy of species named
by Linnaeus [Linné] and other early authors present partic-
ular problems, since many of them named species not on
the b
asis material they personally examined but by reference
to earlier literature sources. For example, in his Systema
naturae, Linnaeus (1758) based most species names on ear-
lier accounts by himself and others. These have been traced
when possible and the exact bibliographic references given.
These secondary references have been examined too; very
often this actually creates additional problems because these
secondary authors themselves refer to older publications, etc.
I have not always included such secondary references and
have only rarely searched the tertiary and earlier sources.
Additionally, several of these earlier works exist in various
editions and Linnaeus' (and other's) bibliographic references
are not detailed enough to decide which editions were used.
For example, I have had the rare opportunity of examining
side by side different editions of Gesner's Nomenclator aqua-
tilium and Fischbuch but could not find all texts to which
Linnaeus referred. As these books usually are considered to
be antiquities or collector-items, interlibrary loans or photo-
copies are not possible. Examination and comparison of the
various editions would mean travelling to a number of li-
braries and investing a lot of time and money, beyond the
limits of the present work. Although there is an obvious his-
torical and academic interest, the utility of the exercise is not
obvious in the context of biological research and usable out-
puts. In such instances, I merely list the reference as given
by 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) on
which the account is based. Where holotypes are extant, or if
lectotypes or neotypes have been designated, the locality of
these specimens of course becomes the type localities. To
identify the type specimens of nominal species described by
these earlier authors, one has to follow about the same pro-
cedure, that is, to find the specimens on which the accounts
cited 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 search
some 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 Linnaeus
are not sacred, however, since Systema naturae has been
designated by later taxonomists as the starting point of to-
day's nomenclatural system, the names he used are the first
available names for the concerned taxa. By the simple logic
of the principle of priority they will remain, regardless of
how usable or informative the descriptions are—and they
usually are useless without recourse to other sources).
Incertae sedis, genera inquirenda, species inquirendae,
nomina dubia. — Incertae sedis are valid family, genera
and species of uncertain taxonomic position. Genera incer-
tae sedis are listed at the beginning of the Order or Family
to which they belong. For example, Pimelodus javus clear-
ly is a member of Siluriformes, but cannot be placed in any
family; it is therefore listed under Siluriformes, before the
family accounts.
Species incertae sedis are listed at the beginning of the fam-
ily to which they belong. Sometimes they are placed in the
genus in which my experience or that of knowledgeable
colleagues suggests they may belong. Alternatively, for spe-
cies placed in genera to which there is a suspicion they do
not belong, the generic name is placed in single quotation
marks (e.g. 'Genus' species), sometimes with a comment
under Taxonomic notes. The fate of a species incertae sedis
is 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 allow
a decision as to whether or not the species is valid. Such
names are listed immediately under the heading of the ge-
nus to which they belong. Species inquirendae that cannot
be 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 in
front 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 original
description 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 the
last 15 years falls into the category species inquirendae.
The fate of a species incertae sedis is that future studies will
show to which genus or family they belong. The fate of a
species inquirenda is to be redescribed and either found to
be a valid species or a synonym of some other species.
A species inquirenda should not be confused with a nomen
dubium. A nomen dubium (plural: nomina dubia) is a name
of doubtful application that is impossible to link with a
known species, or that may apply to several species. Typi-
cally, a nomen dubium would have been described in the
18th 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 dried
specimens brought to Europe by seamen in the 18th centu-
ry; or many species described from Chinese paintings in the
19th century: these paintings usually were not based on a
given specimen but often were an artistic or idealised view
of the species, copied from earlier classical paintings, or
sometimes simply imaginary). The fate of a nomen dubium
is not to remain so, but to become either a valid name or a
synonym after either taxonomic examination or appropriate
nomenclatural decisions.
A genus inquirendum (plural: genera inquirenda) is a ge-
neric name that can be placed in a family but whose de-
Kottelat: Inland fishes of Southeast Asia
scription and associated species (usually species inquirendae
or nomina dubia) do not allow a decision as to whether or
not it is valid. Such names are listed immediately under the
headings of the family to which they belong.
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 seems
necessary to start with a long introduction explaining basic
principles of nomenclature, terminology, and how to under-
stand the data in the present list. More experienced readers
will probably not need to read this section.
The most basic principle of nomenclature is that it deals
with only the names of organisms not with the organisms
themselves. The confusion between animals and their names
mars many taxonomic discussions and is becoming increas-
ingly common and damaging. It is of concern that even the
editors of some scientific journals are no longer able to make
this distinction, especially in fashionable areas like molecu-
lar systematics.
Nomenclature is about the correct formation and treatment
of names and the objective application of the 'legal' criteria
of a code, irrespective of taxonomic concepts or philosoph-
ical approaches. Taxonomy is about the scientific study of
organisms and includes a level of subjective interpretation
of observations that may differ among scientists.
Code. — Here, the word Code refers to the International
Code of Zoological Nomenclature. The current (4th) edi-
tion was published in 1999 and superseded the previous
editions with effect from 1 January 2000. The Code is pub-
lished under the responsibility of the International Com-
mission of Zoological Nomenclature (ICZN,,
a body of zoologists (29 as of 2010), independent of politi-
cal or national entities. Under exceptional circumstances and
following a prescribed procedure, the ICZN has the power
to 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 follow
the Code. Treating a name as a subjective synonym is a tax-
onomic act, not a nomenclatural act.
Original descriptions. — Original description within the
meaning 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 valid
species. 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 not
automatically a valid name.
Since 2012, the Code allows the publication of new names
in electronic-only publications if they fulfill a number of
conditions. Among them, the work must have an ISSN or
ISBN number, be archived, and be registered in Zoobank
Valid name. — A valid name is the correct name applied to
a species. To be valid, a name must first be available. But an
available name is not automatically valid (junior synonyms
are available names but invalid). A 'valid name' should not
be confused with a 'valid species'.
A nominal species is any of the available names created for
a species, irrespective of its validity. If a valid species has x
synonyms, the valid name and the x synonyms are x+1 nom-
inal species.
Species-group names, genus-group names, family-group
names. — The species-group includes all the names of taxa
of 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 ranked
above the genus-group: superfamily, family, subfamily,
tribes, etc.
Spellings. — A basic principle of nomenclature is that the
original 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 spellings
must be corrected (incorrect original spelling). There are
no spelling that may be corrected, there are only spellings
that must be corrected or that must not be corrected. The
main categories of corrections is that if the species name is
a Latin adjective it must agree in grammatical gender with
the gender of the genus name. Incorrect original spellings
should never be used. Corrections of incorrect original spell-
ings allowed by the Code are called justified emendations.
Any intentional (explained) corrections not allowed by the
Code is an unjustified emendation and should never be used;
unjustified emendations are available names with their own
author and date, are objective synonyms of the emended
names, may be homonyms and may be used as substitute
names. They are included in the list but I may have over-
looked some. Any unexplained change or error is called an
incorrect subsequent spelling. Incorrect subsequent spell-
ings are not mentioned in synonymies, except if they have
used 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 the
publication date is more or less equivalent to the date of
printing, in the context of nomenclature, the Code defines
that the date of publication is the date at which a work could
first be obtained (for sale or by free distribution). The date
of publication of a work is important to determine priority
between two synonyms, two homonyms or two nomencla-
tural acts. The Code rules that the earliest one has priority
over the youngest one (senior synonym vs. junior synonym;
senior homonym vs. junior homonym). Even a difference of
one day is enough to give priority. The Code art. 23.9 al-
lows exceptions (reversal of precedence), but only if very
precise conditions are met.
The year of publication is retained as printed on the publi-
cation. It may happen, however, that the work appeared at a
date different from that printed on the publication. If a dif-
ferent date is documented by reliable information, this date
must be retained for nomenclatural purposes. I have not at-
tempted to check the effective publication date of all cited
works; this would have been very time and effort-consum-
ing. I have invested time for such search only when it was
necessary 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 the
Code, or if there was a suspicion that a stated date is far
from the actual date. Otherwise, I consider that usage of the
date printed on the publication is more important than the
actual publication date (e.g. for retrieval on library shelves
or interlibrary loans).
Problems relating to dates associated with taxa are discussed
under Nomenclatural Notes. Those associated with specific
publications are mentioned in the Literature Cited section,
under the respective titles. Some of the more complex cases
are discussed separately: see Bibliographic Notes.
If nomenclatural acts are available from a valid electronic-
only publication, the date of publication is the date of first
distribution. New names and nomenclatural acts first made
available in electronic publications as 'accepted manuscript'
or 'uncorrect proofs' are not available. Taxonomists should
not circulate manuscripts or proofs because this is a serious
source 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 published
before another one. Precedence indicates that a name must
be 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 followed
by the citation of their 'author'. For example the name of the
carp is Cyprinus carpio Linnaeus, 1758, in which Linnaeus
is the author of the name C. carpio and 1758 the date of the
original description. The citation of the author and year is
merely a bibliographic reference. It is in no case an indica-
tion of ownership. Unfortunately, many scientists forget or
ignore the purpose of the mention of the author's name and
this has sometimes resulted in unjustified emotional reac-
tions of authors when one of 'their' species is treated as an
invalid synonym by others. Somehow, naming a new spe-
cies is the formulation of an hypothesis and there is no shame
if an hypothesis is found to be erroneous. Vexed ego may
never accept a synonymy.
The name of an author is written in parentheses when the
name is moved to another genus by subsequent authors. For
example, Ompok bimaculatus (Bloch, 1794) was originally
described as Silurus bimaculatus by Bloch (1794).
For nomenclatural purposes, the Code (art. 50) defines the
author(s) of a work as the person(s) who first published a
name in a way that makes it available. In most cases the
author of a name is the person whose name appears as the
author of the book or article. In some cases of works by
more than one author, if one author only is responsible for
the name, then that person is author of the name and the
name is then cited in the format Barbus binotatus Valenci-
ennes, in Cuvier & Valenciennes, 1842. I personally think
that 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 it
seems even less desirable since the authors of the work had
themselves decided that the work would appear under both
names. In my eyes it serves no nomenclatural purpose to
search who did what in such a work (of course I understand
the historical interest). But the Code says so and even the
ICZN has ruled so on this precise case.
If it is established that a person other than the person named
as the author(s) of a work is alone responsible for both the
name and the conditions making the name available, then
that person is the author of the name (often called second-
ary author). For example, in Schneider (1801), the name
and the description of Mugil cirrhostomus are from unpub-
lished notes of Forster. Schneider had no specimen and no
other source to describe the species; therefore Forster is
author of the name, and the name is cited as M. cirrhosto-
mus Forster, in Schneider, 1801. [Forster's manuscript was
later published by Lichtenstein (1844) and the two texts can
be compared.]
If an external person is author of the description (for exam-
ple personal notes) and the named author of the work cites
him 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 but
choose other names (for example to avoid homonymy); in
this case, Forster is not responsible for both the name and
the conditions making it available, and therefore Schneider
alone is the author.
If an external person merely suggested a name for a species
described in a work, this does not make him the author of
the 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. For
example, when Valenciennes (1846) described Cobitis fas-
ciata he commented that in their notes the collectors (Kuhl
Kottelat: Inland fishes of Southeast Asia
and van Hasselt) had a drawing of this fish, which they had
labelled 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 new
species and wrote the description himself. This makes him
alone the author. The species must be cited as C. fasciata
Valenciennes, in Cuvier & Valenciennes, 1846 and not C.
fasciata Kuhl & van Hasselt, in Cuvier & Valenciennes, 1846.
Posthumous works belong to these categories but can be
more complex. Sometimes it is simply a matter of some-
body publishing a completed manuscript by a deceased
friend 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 means
that if both the names and the descriptions of the new taxa
are entirely the work of a deceased person, then he is the
author. To organise the actual publication is excluded from
the conditions of authorship.
If the description of a given species has been modified by
the editor, both original writer and editor might be co-au-
thor of the name. If the description is completely rewritten
by the editor, based on his own observations, then he is the
author of the name.
Examples: Forsskål died during a travel around the Red Sea.
His notes were later assembled, organised and published by
Niebuhr, using the names in Forsskål's notes and without
work on the content of the text. Forsskål is author of the
names. 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 he
collected in Brazil. He had supervised the preparation of
most plates and had named the species on the plates. Agas-
siz was hired to write a text that could be distributed with
the plates. He wrote the descriptions of all species. For some
he ignored the names on the plates and created new names,
and he is author of these names. For the species names that
appear only in plates, created by Spix, because the plates
alone are among the conditions sufficient to make a name
available, Spix is the author. And in the cases Agassiz wrote
descriptions and used the names created by Spix on the
plates, 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 whose
name determines the validity of a genus. If several species
are placed in genus X, with type species Xx and this genus
is later divided into two genera, the genus which include
species Xx will continue to be genus X while the other ge-
nus will have another name. If two genera have the same
type species, they are objective synonyms. Genus-group
names proposed after 1930 without the fixation of a type
species are not available (Code art. 13.3) [note that art. 13.3
requires that the fixation be "in the original publication [Art.
68]" and that art. 68 includes, as "type species fixed in the
original publication", those established by original designa-
tion, by monotypy, by absolute tautonymy and by Linnean
Type genus. — Each family-group name has a type genus.
The type genus is the genus whose name has been used to
form the name of the family. For example Silurus is the name
of the genus used to form the family group names Siluridae,
Siluriformes, Silurinae, etc. For nomenclature purposes these
three words are a single name. Whatever the rank within the
family-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 the
name 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. It
is 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. The
type 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 which
species the name must be applied. If the type specimen of
the name Yus belongs to species 1, then the name of species
1 is Yus. If the type specimens of the names Yus and Xus
belong 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 are
paratypes and paralectotypes but have no nomenclatural
function. Other 'type' categories (e.g. allotypes, topotypes,
paratopotypes, paraneotypes) are not recognised by the
Code, should not be used and are ignored here. Among them,
allotype is sometime used to designate one of the paratypes
of a sex different from that of the holotype; topotype is used
as 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 author
of work conditions itself of name is
is making name created by
available is
A B B B, in A
A A and B B A & B, in A
A A and B A A & B, in A
to the author, if there is clear evidence that the author based
the nominal species on a single specimen. There is only one
holotype per species. In all cases where there is clear evi-
dence that the author based the nominal species on more
than one specimen (including literature records) but did not
designate 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 specimens
are 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. In
some 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; it
then has the same value as the holotype. The remaining syn-
types then become paralectotypes and lose their status as
primary types. Lectotypes are designated when it is demon-
strated or suspected that the type series includes more than
one species; it allows the name to be definitively fixed to
the nominal species to which the lectotype belongs. Inci-
dentally, the designation of a lectotype also restricts the type
locality to the locality of the lectotype, excluding the local-
ities of the other syntypes. Paratypes and paralectotypes are
not 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 not
been preserved, are lost, or destroyed) and if the name can-
not be unambiguously linked to a valid species, then (and
only then) a specimen can be designated as neotype that
will have the same function as the holotype. Incidentally,
the designation of a neotype also restricts the type locality
to the locality of the neotype. All designations of neotypes
that do not fully satisfy these and several other conditions
laid down in the Code are invalid and must be ignored.
A number of neotype designations are invalid because the
need for a neotype is not stated or demonstrated. This re-
quirement did not exist in the 1985 Code (art. 75(b)) and an
implicit justification was enough. The requirement became
explicit in the 1999 Code, with an added clause (art. 75.3.1)
requiring "a statement that [the neotype] is designated with
the express purpose of clarifying the taxonomic status or
the type locality of a nominal taxon". Unfortunately the 1999
Code is missing a clause explaining what should be done of
neotype designation validly made before 2000 but invalid
under the 1999 Code because of the absence of the state-
ment. Formally, they become invalid because art. 86.3 states
that all former editions of the Code have no force.
This I interpret as an oversight of the editors of the 1999
Code and is unintentional, otherwise it could be the cause
of very serious instability in some groups, which would be
totally against the spirit of the Code. Many of the pre-2000
neotype designations do not have the statement required by
the 1999 Code. I have retained as valid the neotypes validly
designated before 2000 under the 1985 Code.
After the original description, it may be necessary for later
authors to re-examine the primary type of a nominal species
in order to decide to which taxonomic species it applies, for
example in cases when several similar species are later dis-
covered and the original description does not mention the
characters 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 should
not be permitted to handle them. Primary types must be
deposited in museums or other responsible institutions and
with staff able to conserve them and make them accessible
to later researchers. Even if there is political pressure in some
countries to consider types as national property, types do
not 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 recognised
institution (e.g. museum).
A number of species described by earlier authors do not have
known 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 later
eaten 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, when
known, the number of specimens in square brackets (exam-
ple: AAA 1234 [2], BBB 1233 [1]). When the primary types
were 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 point
for further search. The source for the catalogue number is
given when it is not the original description; besides, many
of those listed in the original descriptions have also been
checked in published catalogues or in the institutions them-
selves. When there is a series of syntypes, I listed those
I could trace in the literature, but made no effort to trace the
whole series; this would have been tedious, many of them
having 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 the
type status of the specimen is not certain.
Institutional abbreviations used in the text are listed below.
For institutions for which no abbreviations have been used
in literature, the abbreviations follow current use by work-
Kottelat: Inland fishes of Southeast Asia
ers at these institutions (where possible) or Leviton et al.
(1985; Leviton & Gibbs, 1988) or Eschmeyer (2010). I did
not automatically follow Leviton et al. and Eschmeyer as a
standard because for non-US collections the abbreviations
they list are often not those actually used by the institutions
themselves. In case the abbreviations used in these lists dif-
fer from those used by workers at these institutions, I retain
the 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 meaning
it has in the Code, that is a new name applied to a species
that already had a name. Names erroneously used for a spe-
cies other than the one originally described under that name
are misidentifications. Misidentifications are not synonyms
and are not included in this catalogue. The Code rules that
in case an author thinks that two names actually refer to a
single species (i.e., they are synonyms), the name published
first (senior synonym) is the valid name; the name published
later (junior synonym) is invalid (cannot be used). The jun-
ior synonym nevertheless remain available; should a later
author find that the type specimens of the two names actual-
ly refer to different species, the junior synonym might be
used 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 names
are 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 taxonomic
judgement of an author and other authors may disagree; in
the 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 called
homonyms. The Code rules that the name published first
(senior homonym) is the valid name; the name published
later (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 same
genus 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 called
secondary homonyms. Example: Crayracion fluviatilis var.
ocellata Steindachner, 1870 was not a homonym of Tetra-
odon ocellatus Linnaeus, 1758 when established. Later (in
1975), it was treated as a valid species of Tetraodon and its
name became T. ocellatus (Steindachner, 1870), a junior
secondary homonym of T. ocellatus Linnaeus, 1758.
Replacement names. — If a species name becomes invalid
because 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. If
there is no available synonym, then a new replacement name
is established. The new replacement name takes the same
type as the replaced name. In the example above, Dekkers
(1975) treated Crayracion ocellatus as a valid species of
Tetraodon and made it a junior homonym of Tetraodon ocel-
latus. He created the new replacement name T. steindachneri
to replace the junior homonym.
A junior secondary homonym rejected and replaced before
1961 is definitively invalid (there may be exceptions; Code
art. 59.3). But a junior secondary homonym rejected after
1960 but later considered to be in a genus different from the
senior homonym becomes valid again (Code art. 59.4). In
the above example,when Crayracion ocellatus Steindachner,
1870 was treated as a valid species of Tetraodon it had to be
replaced by Tetraodon steindachneri Dekker, 1975. But
T. steindachneri is now considered to be a valid species of
Dichotomyctere and the senior synonym must be reinstated
and the valid name is now D. ocellatus (Steindachner, 1870),
not D. steindachneri.
Occasionally, some authors have replaced names because
they overlooked an already available name that should have
been used as substitute name, or because they did not like
an existing name, or because they found it inappropriate, or
to 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 the
new replacement name Tetraodon steindachneri Dekkers,
1975 it was discovered that the misidentified T. biocellatus
Tirant, 1885 in fact was also a junior synonym of T. ocellatus
(Steindachner, 1870). Therefore T. biocellatus became the
valid substitute name for T. ocellatus (Steindachner, 1870)
and T. steindachneri became a junior synonym of
T. 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 are
names originally intended for categories below the subspe-
cies level (for example: varieties, natios). These names are
not recognised as valid by the Code. They are nevertheless
listed here. For infrasubspecific names, I have indicated the
locality stated by the original author, but have not listed
material. 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 found
an infrasubspecific name validated this way I noted it; if
such subsequent uses are not listed, it does not mean there
is none, just that I did not find one. Most of these valida-
tions have been accidental (the authors were not aware of
the nomenclatural implications of their use of the name) and
appeared in non-taxonomic publications and often escaped
indexing. A search through the whole ichthyological litera-
ture for such accidental validations is not feasible and their
discovery is usually accidental.
An additional problem with the search for infrasubspecific
names and their accidental validations is that access to the
literature of some countries is limited or restricted, and it is
not a coincidence that such actions and biological concepts
were most frequently used in these very countries.
Ending and spelling of species names. — The following
discussions refers only to scientific names, which are writ-
ten in a language supposed to be Latin. When the system of
binominal nomenclature was created, the genus was the
important entity and the species was of secondary signifi-
cance. As a result, the names of the species were made of a
noun (the genus name) and a qualifying word (the species
name), in most cases an adjective or a noun in the genitive.
As genus names are nouns, they have a grammatical gender
(masculine, feminine or neuter). In Latin, French, German
and most western languages the ending of adjectives varies
to agree with the gender of the noun (English is a notable
exception in which adjectives are not variable). When new
research shows that a species must be transferred from one
genus to another, if the two genera have names with differ-
ent grammatical genders, it may happen that the ending of
the name of the species must be changed to agree in gender
with the name of the genus (Code art. 31.2).
This may seem complicated, but actually it does not require
to learn Latin, but only to follow a handful of trivial rules:
a) check the gender of the name of the genus in the original
publication or on a reliable list;
b) if the species name is a noun, its spelling remains un-
c) if the species name is an adjective, the name must agree
in gender with the genus.
I provided elsewhere (Kottelat, 2012b: 8–12) guidelines on
how existing names should be analysed (Note, this is for
existing names not for creating new names). These guide-
lines apply to the vast majority of names; a few rare cases
make exceptions and are not discussed. Besides, answers to
most questions can be found in dictionaries and in case of
doubt, it does not take a great effort to ask knowledgeable
colleagues. In the last resort, if nobody knows or if it seems
too time consuming, one should treat the name as a noun in
apposition and simply retain the original spelling.
e.g., exempli gratia (for example)
i.a., inter alia (among other things)
I., island
Is., islands
masl, meters above sea level
q.v., quod vide (which see, see there)
s.l., sensu lato (in the wider sense)
s.s., s.str., sensu stricto (in the stricter sense)
viz., videlicet (that is, namely)
Ichthyological collections
AFAQ Museum of Amateur Fishermen's Association of
Queensland, Australia [now in QM]
AMNH American Museum of Natural History, New
York, USA
AMS Australian Museum, Sydney, Australia
ANSP Academy of Natural Sciences, Philadelphia, USA
ASIO Institute of Oceanography, Academia Sinica,
Qingdao, China
ASIZB Institute of Zoology, Academia Sinica, Beijing,
ASIZP Biodiversity Research Museum, Academia Sini-
ca, Taipei, Taiwan
BDSSI Laboratory of the Biology Department, Shang-
hai Science Institute, Shanghai, China [present
status unknown]
BLG Biological Laboratory, Sun Yat-Sen University,
Guangdong, China
BMNH Natural History Museum [formerly British Mu-
seum, Natural History], London, U.K.
BOC Bingham Oceanographic Collection, Yale Uni-
versity, New Haven, USA [now YPM]
BPBM Bernice P. Bishop Museum, Honolulu, USA
BSM Bureau of Science, Manila, Philippines [de-
stroyed during World War II]
CAS California Academy of Sciences, San Francisco,
CM Carnegie Museum [now in FMNH]
CSIRO Australian National Fish Collection, CSIRO,
Hobart, Tasmania, Australia
CUMZ Chulalongkorn University Museum of Zoology,
Bangkok, Thailand
DHFRI Dong Hai Fisheries Research Institute, Shang-
hai, China
DHMB Department of Harbours and Marine, Brisbane,
Australia [now at QM]
DVZUT Department of Vertebrate Zoology, University of
Tông-Hop, Hanoi, Vietnam
ECSFI East China Sea Fisheries Research Institute,
Shanghai, China
FAKU Faculty of Agriculture, Kyoto University, Kyo-
to, Japan
FESC Fisheries Experimental Station, Guangdong,
China [now Pearl River Fishery Research Insti-
tute, Chinese Academy of Fishery Sciences,
Guangzhou, China]
FMNH Field Museum of Natural History, Chicago, USA
Kottelat: Inland fishes of Southeast Asia
FRLM Fisheries Research Laboratory, Mie University,
Mie, Japan
GCM Department of Zoology, Government College
University, Lahore, Pakistan
GUZ Gauhati University, Gauhati, India
HNUE Department of Zoology, Faculty of Biology and
Agricultural Technology, Hanoi National Univer-
sity of Education [also Hanoi University of Ped-
agogy], Hanoi, Vietnam
IBSD Institute of Bioresources and Sustainable Devel-
opment, Takyelpat, India
IHB Institute of Hydrobiology, Wuhan, China
ION Museum of Marine Biodiversity, Institute of
Oceanography, Nhatrang, Vietnam
IPMB Universiti Malaysia Sabah, Kota Kinabalu, Sa-
bah, Malaysia
IPPS Fisheries Research Institute Sarawak, Kuching,
IRSM Institut de la Recherche Scientifique de Mada-
gascar, Antananarivo, Madagascar [mostly now
in MNHN]
IRSNB Institut Royal des Sciences Naturelles, Bruxelles,
ISBB Institutul Stiinte Biologice, Bucuresti, Romania
IZA Dipartimento di Scienze Ambientali, Universi-
ta, L'Aquila, Italy
IZPAN Zoology Institute, Polish Academy of Sciences,
Warszawa, Poland
JNH Department of Biology, Jinan University, Guang-
zhou, China
KIZ Kunming Institute of Zoology, Kunming, China
LSL Linnean Society, London, U.K.
LUG Lingnan University, Guangzhou, China [present
status unknown]
MAMU Macleay Museum, University of Sydney, Syd-
ney, Australia
MARNM Maejo Aquatic Resources Natural Museum,
Chiang Mai, Thailand
MCSNG Museo Civico di Storia Naturale, Genova, Italy
MCZ Museum of Comparative Zoölogy, Cambridge,
MCZL Musée Cantonal de Zoologie, Lausanne, Swit-
MFLB Marine Fisheries Laboratory, Department of
Fisheries, Bangkok, Thailand
MGAB Museul de Istorie Naturala 'Gr. Antipa', Bucuresti,
MGHNL Musée Guimet d'Histoire Naturelle, Lyon, France
MHNG Muséum d'Histoire Naturelle, Genève, Switzer-
MHNN Musée d'Histoire Naturelle, Neuchâtel, Switzer-
MIKU Marine Biological Institute, Kyoto University,
Japan [now at FAKU]
MMNHN Metropolitan Museum of Natural History, Nank-
ing, China [now Nanjing Museum, Chinese
Academy of Sciences, Nanjing, China]
MNCN Museo Nacional de Ciencias Naturales, Madrid,
MNH Magyar Nemzeti Múzeum, Budapest, Hungary
MNHN Muséum National d'Histoire Naturelle, Paris,
MSNM Museo Civico di Storia Naturale, Milano, Italy
MTD Museum für Tierkunde, Dresden, Germany
MUMF Manipur University Museum of Fishes, Can-
chipur, India
Muzium Sabah, Kota Kinabalu, Sabah, Malaysia
MZB Museum Zoologicum Bogoriense, Cibinong, In-
MZUB Museo Zoologico dell'Università de Bologna,
Bologna, Italy
MZUF Museo Zoologico 'La Specola', Università di
Firenze, Firenze, Italy
MZUSP Museu de Zoologia, Universidade de São Paulo,
São Paulo, Brazil
MZUT Museo Zoologico, Universita, Torino, Italy
NCNTTSI Research Institute for Aquaculture No. 1 [Vien
Nghien curu Nuoi trong Thuy san 1; earlier Dinh
Bang Fish Research Station], Bac Ninh, Vietnam
NHMG Naturhistoriska Museum, Göteborg, Sweden
NIG Nanjing Institute of Geography and Limnology,
Chinese Academy of Sciences, Nanjing, China
NKMC National Kweiyang [Guiyang] Medical College,
Guiyang, Guizhou, China [present status un-
NMBA Naturhistorisches Museum, Basel, Switzerland
NMBE Naturhistorisches Museum, Bern, Switzerland
NMMB National Museum of Marine Biology and Aquar-
ium, Pingtung, Taiwan
NMSL National Museum of Sri Lanka, Colombo, Sri
NMSZ National Museums of Scotland, Edinburgh, Scot-
land, U.K.
NMV Museum Victoria, Melbourne, Australia
NMW Naturhistorisches Museum, Wien, Austria
NPIB Northwest Plateau Institute of Biology, Chinese
Academy of Sciences, Xining, Qinghai, China
NRIBAS National Research Institute Biology, Academy
of Sciences, Nanjing, China
NRM Naturhistoriska Riksmuseet, Stockholm, Sweden
NSMT National Science Museum, Tokyo, Japan
NTM Museum and Art Gallery of the Northern Terri-
tory, Darwin, Australia
NTUM National Taiwan University, Taipei, Taiwan
NZOI New Zealand Oceanographic Institute [now Na-
tional Center for Coasts and Oceans], Welling-
ton, New Zealand;
OUC Ocean University of China, Qingdao, China
PNM Philippines National Museum, Manila, Philip-
QM Queensland Museum, Brisbane, Australia
RCMMF Museum of Fishes, Research Center Manipur,
Imphal, India
RMNH Naturalis Biodiversity Center [earlier Nationaal
Natuurhistorisch Museum, earlier Rijksmuseum
van Natuurlijke Historie], Leiden, The Netherlands
ROM Royal Ontario Museum, Toronto, Canada
RUSI J. L. B. Smith Institute of Ichthyology, Graham-
stown, South Africa [now SAIAB]
SAIAB South African Institute of Aquatic Biodiversity,
Grahamstown, South Africa
South African Museum, Cape Town, South Africa
SAMA South Australian Museum, Adelaide, Australia
SBC Sarawak Biodiversity Center, Kuching, Sarawak,
SBM Sabah Museum, Kota Kinabalu, Sabah, Malay-
sia [see MUS]
SCNU South China Normal University, College of Life
Science, Department of Biology, Guangzhou,
SCSFRI South China Sea Fisheries Research Institute,
Guangzhau, China
SFC Shanghai Fisheries College, Shanghai, China
[now SFU]
SFI Shanghai Fisheries Institute, Shanghai, China
[now SFU]
SFU Shanghai Fisheries University, Shanghai, China
SMF Forschungsinstitut Senckenberg, Frankfurt, Ger-
SMK Sarawak Museum, Kuching, Sarawak, Malaysia
Slovenské Národné Múseum, Bratislava, Slovakia
SMNS Staatliches Museum für Naturkunde, Stuttgart,
Museum of the Biological Laboratory, Science
Society of China, Nanking, China [now NRIBAS]
SU Stanford University [now at CAS]
SWFC Museum of Zoology, Southwest Forestry Col-
lege, Kunming, China
TFRI Taiwan Fisheries Research Institute, Chilung,
THNHM Thailand Natural History Museum, National Sci-
ence Museum, Pathum Tani, Thailand
TINRO Museum Tinro, Vladyvostok, Russia
TISTR Thai Institute of Science and Technology,
Bangkok, Thailand
TMBU Museum of the Department of Zoology, Tilka-
manjhi Bhagalpur University, Bhagalpur, India
TUF Laboratory of Fishery Biology, Tokyo Universi-
ty of Fisheries [Museum of Fishery Sciences,
Tokyo University of Marine Sciences and Tech-
nology], Tokyo, Japan
TUK Department of Zoology, Tribhuvan University,
Kathmandu, Nepal
Universiti Brunei Darussalam, Brunei Darussalam
UF Florida Museum of Natural History, University
of Florida, Gainesville, USA
UCDZ Fisheries Laboratory, Department of Zoology,
University of Calcutta, Kolkata, India
UMB Ueberseemuseum, Bremen, Germany
UMMZ University of Michigan Museum of Zoology, Ann
Arbor, USA
UMSB University Malaysia Sabah, Kota Kinabalu, Sa-
bah, Malaysia
UMZC University Museum of Zoology, Cambridge,
UNMF Ubonratchathani University Natural History
Museum of Fisheries, Ubonratchathani, Thailand
URM Department of Marine Sciences, University of
the Ryukyus, Japan
USNM National Museum of Natural History, Washing-
ton, USA
UUZM Evolutions-Museet, Uppsala Universitet, Uppsa-
la, Sweden
VUP Vinh University of Pedagogy, Vinh, Vietnam
WAM Western Australian Museum, Perth
WIAP Wistar Institute of Anatomy, Philadelphia, USA
[now at ANSP]
WURC Walailak University Reference Collection, Na-
khon Si Thammarat, Thailand
YCM Yokosuka City Museum, Yokosuka, Japan
YPM Yale University, Peabody Museum, New Haven,
YU School of Life Science, Yunnan University, Kun-
ming, China
ZFMK Zoologisches Forschungsinstitut und Museum
Alexander Koenig, Bonn, Germany
ZISP Zoological Institute of the Academy of Scien-
ces, St. Petersburg, Russia
ZIU Zoological Museum, University of Uppsala,
Sweden [now UUZM];
ZMA Zoölogisch Museum, Universiteit van Amster-
dam, Amsterdam, The Netherlands [now in
ZMAU Zoology Museum, Andhra University, Waltair,
Vishakhapatnam, India
ZMB Museum für Naturkunde, Berlin, Germany
ZMFMIB Zoologial Museum, Fan Memorial Institute of
Biology, Tsing Hua University, Beijing, China
[now ASIZB]
ZMH Zoologisches Museum und Zoologisches Insti-
tut, Hamburg, Germany
ZMMU Zoological Museum, Moscow State University,
Moscow, Russia
ZMUAS Zoological Museum, Academy of Sciences, Kiev,
ZMUC Zoologisk Museum, København, Denmark
ZMUO Universitetets I Oslo, Zoologisk Museum, Oslo,
ZMUR Zoological Museum, University of Rangoon,
ZMUU Zoologiska Museet, Uppsala Universitet, Upp-
sala, Sweden [now UUZM]
ZMZ Zoologisches Museum, Zürich, Switzerland
ZRC Raffles Museum of Biodiversity Research, Na-
tional University of Singapore, Singapore
ZSI Zoological Survey of India, Calcutta, India
ZSI/CRS Central Regional Station, ZSI, Jabalpur, India
ZSI/NRS Northern Regional Station, ZSI, Dehra Dun, In-
ZSI/SRS Southern Regional Station, ZSI, Madras, India
ZSM Zoologische Staatssammlung, München, Germa-
ZUMT Department of Zoology, University Museum,
University of Tokyo, Tokyo, Japan
Kottelat: Inland fishes of Southeast Asia
not Indian Ocean as reported by Gmelin]; holotype:
BMNH 1853.11.12.205, Wheeler, 1958: 203)
Squalus tuberculatus Bloch, in Schneider, 1801: 137 (ap-
parently based only on Squale dentelé of La Cepède, 1798:
281, pl. 11 fig. 1; type locality: unknown; holotype:
MNHN; ZMB 4443 [2, Paepke & Schmidt, 1988: 162]
unlikely to be types)
Squalus Denticulatus Shaw, 1804b: 351 (based only on
Squale dentelé of La Cepède, 1798: 281, pl. 11 fig. 1;
type locality: unknown; holotype: MNHN; objective jun-
ior synonym of Squalus tuberculatus Bloch, in Schneider,
1801: 137)
Squalus Gronovianus Shaw, 1804b: 353 (unnecessary re-
placement name for Squalus indicus Gmelin, 1789: 1503)
? Squalus Variegatus Blainville, 1816: 121 (nomen nudum)
Squalus Dentatus Blainville, 1816: 121 (nomen nudum)
? Squalus Lambarda Blainville, 1816: 121 (nomen nudum)
Chiloscyllium phymatodes Bleeker, 1852a: 21 (type locali-
ty: Indonesia: Java: Samarang; holotype [383 mm TL]:
? RMNH 7406 [1 of 2, smaller specimen], Dingerkus &
DeFino, 1983: 22)
Squalus caudatus Gronow, in Gray, 1854: 8 (type locality:
Indian Sea [East Indies; Gronovius, 1763: 34]; holotype:
BMNH 1853.11.12.205, Wheeler, 1958: 203; objective
junior synonym of Squalus indicus Gmelin, 1789: 1503)
Chiloscyllium colax Whitley, 1939a: 228 (available by indi-
cation to Meuschen, 1781: [3], itself based on "n°150",
which is Squalus dentibus acutis of Gronovius, 1763: 34,
n°150; type locality: East Indies [Gronovius, 1763: 34];
holotype: BMNH 1853.11.12.205, Wheeler, 1958: 203;
objective junior synonym of Squalus indicus Gmelin,
1789: 1503)
Taxonomic notes. Often enters freshwater and brackish ar-
eas (Last et al., 2010: 50). Synonymy based on Dingerkus &
DeFino (1983). Squalius colax Meuschen, 1781 treated as
valid by some authors (e.g. Kharin, 1987: 69) is not an avail-
able name.
Chiloscyllium Müller & Henle, 1837
Chiloscyllium Müller & Henle, 1837a: 112 (type species:
Scyllium plagiosum Bennett, 1830: 694, by subsequent
monotypy in Smith, 1838b: 85; also in Müller & Henle,
1837b: 395, 1838a: 17, 1838b: 34). Gender neuter.
Synchismus Gill, 1862a: 407, 408 (type species: Squalus tu-
berculatus Bloch, in Schneider, 1801: 137, by original
designation; also in Gill, 1862h: 413). Gender mascu-
Taxonomic notes. Revision by Dingerkus & DeFino (1983).
Nomenclatural notes. The original description of Chiloscyl-
lium (Müller & Henle, 1837a: 112) was in a summary of
their forthcoming monograph on plagiostomes (Müller &
Henle, 1838a: 17). The same text also appeared in Müller &
Henle (1837b: 395) and was translated in English (Müller &
Henle, 1838b: 34), both without included species. The first
mention of a species name in combination with Chiloscyl-
lium is by Smith (1838b: 85) in the proceedings of a meeting
held on 12 September 1837, published on 13 February 1838.
Smith listed various genera, commenting that he had just seen
Müller & Henle's work (obviously 1837a or 1837b); he cited
Chiloscyllium, including a single species (C. plagiosum),
making it type species [Incidentally, the article mentions that
Müller was present at the meeting]. The name was then next
used by Müller & Henle (1838c: 83), without included spe-
cies. In this article they announced that the first parts of 1838a
will still appear the same year. Article 1838c cannot be dated,
but the previous page in the volume (p. 82) mentions a meet-
ing held in London on 5 December 1837.
Chiloscyllium indicum (Gmelin, 1789)
[Squalus] colax Meuschen, 1781: [3] (not available, pub-
lished in a rejected work; ICZN, 1954e: 281, Opinion
Squalus indicus Gmelin, 1789: 1503 (based on Squalus den-
tibus acutis of Gronovius, 1754: 61, n°133, 1763: 34,
n°150; type locality: East Indies [Gronovius, 1763: 34;
Taxonomic notes. For phylogeny and classification, see Goto
Stegostoma Müller & Henle, 1837
Stegostoma Müller & Henle, 1837a: 112 (type species:
Squalus fasciatus Bloch, 1784b: 19, by original designa-
tion; also in Müller & Henle, 1837b: 395, 1838b: 35).
Gender neuter.
Stegostoma tigrinum (Forster, 1781)
Squalus varius Seba, 1759: 105, pl. 34 fig. 1 (not available,
because binominal nomenclature not used consistently;
also applies to Index)
Squalus tigrinus Forster, 1781: 24, pl. 13 fig. 2 (type locali-
ty: Indian Ocean / Sri Lanka [Singhalese name]; syntypes:
Forster's material and 2 specimens of Seba, 1759: pl. 34
fig. 1)
Squalus fasciatus Hermann, 1783: 302, table (based on Seba,
1759: pl. 34 fig. 1; type locality: no data; lectotype: spec-
imen figured in Seba, 1759: pl. 34 fig. 1, designated by
Fricke, 1999a: 13; spelling Squalo fasciato on p. 302, a
non-nominative declension to be corrected in Squalius
fasciatus (Code art. 11.9.2); mention of Linné, 1767: 401
relates to Chimaera not S. fasciatus)
Squalus tigrinus Broussonet, 1784: 659 (based on Squalus
varius Seba, 1759: 105, pl. 34 fig. 1, Gronovius, 1754:
62, n°136 [itself based on Seba, 1759: 105], Forster, 1781:
24, pl. 13 fig. 2; type locality: "mer des Indes" [sea of the
Indies] / China: "rivière de Canton" [river of Canton];
syntypes: material of Seba and Forster; junior primary
homonym of Squalus tigrinus Forster, 1781: 24)
Squalus fasciatus Bloch, 1784b: 19, pl. 113 (type locality:
India: Tranquebar [Tharangambadi, 11°01'37"N 79°51'E];
syntypes: ZMB 4449 [1, listed as holotype], 7833 [1],
22610 [1], Paepke & Schmidt, 1988: 163; primary junior
homonym of Squalus fasciatus Hermann, 1783: 302); also
Bloch, in Schneider, 1801: 130 (locality: "Indian Ocean
at the mouth of Coromandel")
Squalus tygrinus Bonnaterre, 1788: 8, pl. 8 fig. 23 (based on
Bloch, 1787b: 17, Forster, 1781: 24, pl. 13 fig. 2, Brous-
sonet, 1784: 658; type locality: "la mer des Indes"; syn-
types: material on which are based the cited references)
Squalus fasciatus Bonnaterre, 1788: 8 (based on 'galloné of
Broussonet, 1784: 659; type locality: South Africa: Cape
of Good Hope; holotype: BMNH)
Squalus tigrinus Gmelin, 1789: 1493 (based on Forster, 1781:
pl. 13 fig. 2, Squalus fasciatus Bloch, 1784b: 19, pl. 4
[113], Gronovius, 1754: 62, n°136 [ex museo Sebae],
1763: 33, n°147, Seba, 1759: 105, pl. 34 fig. 1, Hermann,
1783: 302, Broussonet, 1784: 658; type locality: Indian
Ocean; syntypes: material of these authors; junior prima-
ry homonym of Squalus tigrinus Forster, 1781: 24)
Squalus longicaudus Gmelin, 1789: 1496 (based on
Gronovius, 1754: 62, n°136 [ex museo Sebae], 1763: 33,
n° 147 and Seba, 1759: 105 n° 1, pl. 34 fig. 1; type local-
ity: no data; syntypes [2]: specimens mentioned by Seba)
Squalus tigrinus Pennant, 1791: 92, pl. 13 fig. 2 (type local-
ity: no data; syntypes [2]: specimens mentioned by Seba,
1759: 105; junior primary homonym of Squalus tigrinus
Forster, 1781: 24)
Squalus zebra Shaw, 1804b: 352 (based on Artedi, 1738,
Seba, 1759: pl. 34 fig. 1, Gmelin, 1789, Bloch, 1784b:
pl. 113, type locality: Indian Seas; syntypes: at least ma-
terial mentioned by Seba [2])
Scyllia Quinquecornuatum van Hasselt, 1823a: 315 [trans-
lated in Alfred, 1961b: 81] (type locality: Indonesia: Java;
syntypes: material of Kuhl & van Hasselt, and Seba, 1759:
pl. 34 fig. 1)
Scyllium heptagonum Rüppell, 1837: 61, pl. 17 fig. 1 (type
locality: Red Sea: Saudi Arabia: Didda [Jeddah]; holo-
type: SMF 3152; lectotype designation by Klausewitz,
1960: 290 is invalid as Rüppell explicitly stated having
seen a single specimen and did not give bibliographic
indication to additional sources)
Stegostoma carinatum Blyth, 1847: 725, pl. 25 fig. 1 (type
locality: India; holotype: ? ZSI)
Squalus pantherinus Bleeker, 1852a: 23 (not available, name
listed in synonymy)
Squalus cirrosus Gronow, in Gray, 1854: 6 (based on
Gronovius, 1763: 33, n° 147 [itself based on Gronovius,
1754: 62, n°136, itself on Seba], Seba, 1759: pl. 34 fig.
1; type locality: no data; syntypes [2]: specimens men-
tioned by Seba)
Stegostoma varium Garman, 1913: 59 (type locality: India
and East Indies to Africa; syntypes: MCZ 55-S [1, Phil-
ippines: Manila], ? 33437 [1, Mauritius], uncat. [1], Esch-
meyer, 2010, and material on which cited references are
Stegostoma tigrinum naucum Whitley, 1939a: 229, fig. 2
(type locality: Australia: NSW: Hawkesbury River; ho-
lotype: AMS I.4174, Paxton et al., 1989: 92)
Scyllium quinquecarinatum Compagno, 1984: 200 (errone-
ous subsequent spelling of Scyllia quinquecornuatum van
Hasselt, 1823a: 315)
Taxonomic notes. Freshwater record from Philippines by
Herre (1925c: 126). Synonymy adapted from Compagno
(1984: 200). This species is commonly mentioned as Stego-
stoma fasciatum, but the name S. tigrinum has priority. As
the latter has been used after 1899, it cannot be suppressed
under Code art. 23.9.1.
Kottelat: Inland fishes of Southeast Asia
Carcharhinus Blainville, 1816
Carcharhinus Blainville, 1816: 121 (subgenus of Squalus
Linnaeus, 1758: 233; type species: Carcharhias melan-
opterus Quoy & Gaimard, 1824: 194, designated by
ICZN, 1965: 32 [Opinion 723.2c]; on Official List of
Generic Names in Zoology, ICZN, 1965: 32 [Opinion
723.3a]). Gender masculine.
Carcharias Cuvier, 1816a: 125 (subgenus of Squalus Lin-
naeus, 1758: 233; type species: Squalus carcharias Lin-
naeus, 1758: 235, by absolute tautonymy; junior hom-
onym of Carcharias Rafinesque-Schmaltz, 1810a: 10; on
Official Index of Rejected and Invalid Generic Names in
Zoology, ICZN, 1965: 33 [Opinion 723.5c]). Gender