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An International Journal of Southeast Asian Zoology
Supplement No. 27 22 November 2013
The Raffles Bulletinof Zoology
Published by the Department of Biological Sciences, National
University of Singapore
Articles appearing in this journal are indexed in: SCIENCE
CITATION INDEX® CURRENT CONTENTS® AGRICULTURE, BIOLOGY &
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ABSTRACTS® CAMBRIDGE SCIENTIFIC ABSTRACTS® AQUATIC SCIENCES &
FISHERIES ABSTRACTS
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The Fishes of the Inland Waters of Southeast Asia:
A Catalogue and Core Bibliography of the Fishes Known to Occur
in Freshwaters, Mangroves and Estuaries
Maurice Kottelat
TH
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AFFL
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OF Z
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2013 SU
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SUPPLEMENTS PUBLISHED(continued from inside cover)
7. Allwood, A. J., A. Chinajariyawong, R. A. I. Drew, E. L.
Hamacek, D. L. Hancock, C. Hengsawad, J. C. Jipanin, M. Jirasurat,
C. Kong Krong, S. Kritsaneepaiboon, C. T. S. Leong & S.
Vijaysegaran, 1999. Host plant records for fruit fl ies (Diptera:
Tephritidae) in Southeast Asia. Raffl es Bulletin of Zoology,
Supplement 7: 1–92.
8. Ng, P. K. L. & K. S. Tan (eds.), 2000. The biodiversity
of South China Sea. Raffl es Bulletin of Zoology, Supplement 8:
1–673.
9. Kurahashi, H. & F. R. Magpayo, 2000. Blow fl ies
(Insecta: Diptera: Calliphoridae) of the Philippines. Raffl es
Bulletin of Zoology, Supplement 9: 1–78.
10. Jefferson, T. A. & B. D. Smith (eds.), 2002. Facultative
freshwater cetaceans of Asia: Their ecology and conservation. Raffl
es Bulletin of Zoology, Supplement 10: 1–187.
11. Ng, P. K. L., D. Wowor & D. C. J. Yeo (eds.), 2002.
Scientifi c results of the Anambas Expedition 2002. Raffl es
Bulletin of Zoology, Supplement 11: 1–130.
12. Yeo, D. C. J., P. K. L. Ng & R. Pethiyagoda (eds.),
2005. Contributions to biodiversity exploration and research in Sri
Lanka. Raffl es Bulletin of Zoology, Supplement 12: 1–434.
13. Kottelat, M. & D. C. J. Yeo (eds.), 2005. Southeast
Asian freshwater fi sh diversity. Raffl es Bulletin of Zoology,
Supplement 13: 1–208.
14. Tan, H. & R.-Q. Jan (eds.), 2007. Proceedings of the 7th
Indo-Pacifi c Fish Conference. Raffl es Bulletin of Zoology,
Supplement 14: 1–434.
15. Wang, L. K. & C. J. Hails, 2007. An annotated checklist
of birds of Singapore. Raffl es Bulletin of Zoology, Supplement 15:
1–179.
16. Tan, S. H. & P. K. L. Ng (eds.), 2007. Crustacean
Supplement 1. Raffl es Bulletin of Zoology, Supplement 16:
1–357.
17. Ng, P. K. L., D. Guinot & P. J. F. Davie, 2008. Systema
Brachyurorum: Part 1. An annotated checklist of extant brachyuran
crabs of the world. Raffl es Bulletin of Zoology, Supplement 17:
1–286.
18. Bieler, R., K. Chalermwat, P. M. Mikkelsen, K. S. Tan &
E. Wells (eds.), 2008. Molluscs of Eastern Thailand: Proceedings of
the International Marine Bivalve Workshop, Chantaburi, Thailand,
August–September 2005, with contributions on other molluscan
groups. Raffl es Bulletin of Zoology, Supplement 18: 1–264.
19. Tan, S. H. & I.-S. Chen (eds.), 2008. Aquatic
biodiversity of the South China Sea. Raffl es Bulletin of Zoology,
Supplement 19: 1–292.
20. Tan, S. H. & M. E. Y. Low (eds.), 2009. Crustacean
Supplement II. Raffl es Bulletin of Zoology, Supplement 20:
1–307.
21. De Grave, S., N. D. Pentcheff, S. T. Ahyong, T.-Y. Chan, K.
A. Crandall, P. C. Dworschak, D. L. Felder, R. M. Feldmann, C. H.
J. M. Fransen, L.Y. D. Goulding, R. Lemaitre, M. E. Y. Low, J. W.
Martin, P. K. L. Ng, C. E. Schweitzer, S. H. Tan, D. Tshudy &
R. Wetzer, 2009. A classifi cation of living and fossil genera of
decapod crustaceans. Raffl es Bulletin of Zoology, Supplement 21:
1–109.
22. Tan, K. S. (ed.), 2009. Fourteenth International Marine
Biology Workshop 2006: The marine fl ora and fauna of Singapore.
Raffl es Bulletin of Zoology, Supplement 22: 1–294.
23. Low, M. E. Y. & S. H. Tan (eds.), 2010. Checklists of
anomuran decapod curstaceans of the world (exclusive of the
Kiwaoidea and families Chirostylidae and Galatheidae of the
Galatheoidea) and marine lobsters of the world. Raffl es Bulletin
of Zoology, Supplement 23: 1–181.
24. Davison, G. W. H. & C. S. W. Chia (eds.), 2011.
Proceedings of the fi fth International Hornbill Conference,
Singapore, 22–25 March 2009. Raffl es Bulletin of Zoology,
Supplement 24: 1–176.
25. Koh, L. P., T. M. Lee & M. L. M. Lim (eds.), 2012.
Special Memorial Issue: Navjot S. Sodhi (1962–2011). Raffl es
Bulletin of Zoology, Supplement 25: 1–289.
26. Kottelat, M., 2013. Conspectus cobitidum: An inventory of
the loaches of the world (Teleostei: Cypriniformes: Cobitoidei).
Raffl es Bulletin of Zoology, Supplement 26: 1–199.
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SUPPLEMENTS PUBLISHED
1. Wee, D. P. C. & P. K. L. Ng, 1995. Swimming crabs of the
genera Charybdis De Haan, 1883 and Thalamita Latreille, 1829
(Crustacea: Decapoda: Brachyura: Portunidae) from Peninsular
Malaysia and Singapore. Raffl es Bulletin of Zoology, Supplement 1:
1–128.
2. Deeleman-Reinhold, C. L., 1995. The Ochyroceratidae of the
Indo-Pacifi c region (Araneae). Raffl es Bulletin of Zoology,
Supplement 2: 1–103.
3. Ng, P. K. L. & C. T. N. Chuang, 1996. The Hymenosomatidae
(Brachyura) of Southern Asia, with notes on other species. Raffl es
Bulletin of Zoology, Supplement 3: 1–82.
4. Morioka, H. & C. M. Yang, 1996. A catalogue of the bird
specimens in the Singapore Zoological Reference Collection Part I.
Struthioniformes–Charadriiformes. Raffl es Bulletin of Zoology,
Supplement 4: 1–141.
5. Kurahashi, H., N. Benjaphong & B. Omar, 1997. Blow fl ies
(Insecta: Diptera: Calliphoridae) of Malaysia and Singapore. Raffl
es Bulletin of Zoology, Supplement 5: 1–88.
6. Sodhi, N. S., H. S. Yong & P. K. L. Ng (eds.), 1999. The
biodiversity of Pulau Tioman, Peninsular Malaysia. Raffl es
Bulletin of Zoology, Supplement 6: 1–288.
(continues on back cover)
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THE RAFFLES BULLETIN OF ZOOLOGY 2013
1
THE RAFFLES BULLETIN OF ZOOLOGY 2013 Supplement No. 27: 1–663
http://zoobank.org/urn:lsid:zoobank.org:pub:0B66AE04-C644-43CD-9B76-043848FAA9FE
Date of Publication: 22 Nov.2013© National University of Singapore
and the author
THE FISHES OF THE INLAND WATERS OF SOUTHEAST ASIA :A CATALOGUE
AND CORE BIBLIOGRAPHY OF
THE FISHES KNOWN TO OCCUR IN FRESHWATERS,MANGROVES AND
ESTUARIES
Maurice KottelatCase postale 57, CH-2952 Cornol, Switzerland
(address for correspondence), and
Raffles Museum of Biodiversity Research, Department of
Biological Sciences, National University of Singapore,6 Science
Drive 2, Singapore 117546, Republic of Singapore
Email: [email protected]
CONTENTS
Abstract
.........................................................................
3Introduction
...................................................................
4Basic principles of nomenclature ................................
10Abbreviations used
...................................................... 15
Class ChondrichthyesSubclass Elasmobranchii
Division SelachiiOrder Orectolobiformes
Family Hemiscylliidae .......................... 18Family
Stegostomatidae ........................ 19
Order CarcharhiniformesFamily Carcharhinidae
......................... 20
Division BatoideaOrder Pristiformes
Family Pristidae .................................... 23Order
Rajiformes
Family Rhinobatidae ............................. 24Order
Myliobatiformes
Family Dasyatidae ................................ 25Family
Potamotrygonidae ..................... 28Family Myliobatididae
.......................... 29
Class ActinopterygiiDivision Teleostei
Order OsteoglossiformesFamily Osteoglossidae
............................... 30Family Arapaimidae
................................... 31Family Notopteridae
................................... 31
Order ElopiformesFamily Elopidae
......................................... 33Family Megalopidae
................................... 33
Order AlbuliformesFamily Albulidae
........................................ 34
Order AnguilliformesFamily Anguillidae
..................................... 37Family Moringuidae
................................... 39Family Muraenidae
.................................... 41Family Ophichthidae
.................................. 43Family Muraenesocidae
............................. 48
Order ClupeiformesFamily Pristigastridae
................................. 49Family Engraulididae
................................. 52Family Chirocentridae
................................ 57Family Clupeidae
....................................... 58Family Sundasalangidae
............................. 64
Order GonorhynchiformesFamily Chanidae
......................................... 64
Order CypriniformesFamily Cyprinidae
...................................... 65Family Psilorhynchidae
............................ 171Family Gyrinocheilidae
............................ 173Family Botiidae
........................................ 173Family Vaillantellidae
............................... 176Family Cobitidae
...................................... 176Family Ellopostomatidae
.......................... 184Family Barbuccidae
.................................. 185Family Balitoridae
.................................... 185Family Gastromyzontidae
........................ 191Family Serpenticobitidae
......................... 197Family Nemacheilidae
.............................. 198
Order CharaciformesFamily Characidae
.................................... 215Family Serrasalmidae
............................... 215
Order SiluriformesFamily Loricariidae
.................................. 215Family Amblycipitidae
............................. 216
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Kottelat: Inland fishes of Southeast Asia
2
Family Akysidae .......................................
217Family Sisoridae .......................................
221Family Cranoglanididae ........................... 232Family
Siluridae ....................................... 232Family
Chacidae ....................................... 240Family
Plotosidae ..................................... 240Family
Clariidae ....................................... 242Family Ariidae
.......................................... 245Family Schilbeidae
................................... 251Family Pangasiidae
.................................. 253Family Bagridae
....................................... 256
Order OsmeriformesFamily Plecoglossidae
.............................. 268Family Salangidae
.................................... 268
Order SalmoniformesFamily Salmonidae
................................... 270
Order AulopiformesFamily
Synodontidae................................ 270
Order GadiformesFamily Bregmacerotidae
.......................... 271
Order OphidiiformesFamily Carapidae
..................................... 271Family Bythitidae
..................................... 272
Order BatrachoidiformesFamily Batrachoididae
............................. 272
Order LophiiformesFamily Antennariidae
............................... 274
Order MugiliformesFamily Mugilidae
..................................... 275
Order AtheriniformesFamily Telmatherinidae
............................ 283Family Phallostethidae
............................. 284Family Atherinidae
................................... 286
Order BeloniformesFamily Adrianichthyidae
.......................... 288Family Hemiramphidae
............................ 290Family Zenarchopteridae
......................... 293Family Belonidae
..................................... 297
Order CyprinodontiformesFamily Aplocheilidae
............................... 299Family Poeciliidae
.................................... 300
Order BeryciformesFamily Holocentridae
............................... 301
Order GasterosteiformesFamily Indostomidae
................................ 301Family Syngnathidae
................................ 301
Order SynbranchiformesFamily Synbranchidae
.............................. 307Family Chaudhuriidae
.............................. 309Family Mastacembelidae
.......................... 310
Order ScorpaeniformesFamily Scorpaenidae
................................ 314Family Tetrarogidae
................................. 315Family Synanceidae
................................. 316Family Platycephalidae
............................ 316
Order PerciformesSuborder Percoidei
Family Ambassidae ............................. 318Family
Latidae .................................... 323Family
Lateolabracidae ...................... 324
Family Percichthyidae ........................ 324Family
Serranidae ............................... 325Family
Pseudochromidae ................... 328Family Opistognathidae
...................... 329Family Centrarchidae
.......................... 329Family Apogonidae
............................. 329Family Sillaginidae
............................. 330Family Carangidae
.............................. 331Family Leiognathidae
......................... 337Family Lutjanidae
............................... 340Family Datnioididae
........................... 344Family Lobotidae
................................ 344Family Gerreidae
................................ 345Family Haemulidae
............................. 347Family Nemipteridae
.......................... 349Family Lethrinidae
.............................. 350Family Sparidae
.................................. 352Family Polynemidae
........................... 353Family Sciaenidae
............................... 356Family Mullidae
.................................. 360Family Monodactylidae
...................... 362Family Toxotidae
................................ 362Family Drepaneidae
............................ 365Family Chaetodontidae
....................... 365Family Nandidae
................................. 366Family Pristolepididae
........................ 366Family Badidae
................................... 367Family Terapontidae
........................... 368Family Kuhliidae
................................ 371
Suborder LabroideiFamily Cichlidae
................................. 373Family Pomacentridae
........................ 375Family Labridae
.................................. 379Family Scaridae
.................................. 380
Suborder TrachinoideiFamily Trichonotidae
.......................... 382
Suborder BlennioideiFamily Blenniidae
............................... 382
Suborder CallionymoideiFamily Callionymidae
........................ 385
Suborder GobioideiFamily Rhyacichthyidae .....................
386Family Odontobutidae ........................ 387Family
Eleotrididae ............................ 388Family Kraemeriidae
.......................... 396Family Gobiidae
................................. 396Family Amblyopidae
.......................... 432Family Ptereleotrididae
....................... 435
Suborder KurtoideiFamily Kurtidae
.................................. 436
Suborder AcanthuroideiFamily Ephippidae
.............................. 437Family Scatophagidae
......................... 438Family Siganidae
................................ 439Family Acanthuridae
........................... 441
Suborder ScombroideiFamily Sphyraenidae
.......................... 444Family Scombridae
............................. 446
Suborder AnabantoideiFamily Anabantidae
............................ 446
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THE RAFFLES BULLETIN OF ZOOLOGY 2013
3
Family Helostomatidae ....................... 447Family
Osphronemidae ....................... 448
Suborder ChannoideiFamily Channidae
............................... 458
Order PleuronectiformesFamily Paralichthyidae
............................. 462Family Tephrinectidae
.............................. 463Family Soleidae
........................................ 463Family Cynoglossidae
.............................. 466
Order TetraodontiformesFamily Triacanthidae
................................ 468
ABSTRACT. — There are 3108 valid and named native fish species
in the inland waters ofSoutheast Asia between the Irrawaddy and Red
River drainages, the small coastal drainagesbetween the Red River
and Hainan, the whole Indochinese Peninsula, Andaman and
NicobarIslands, Indonesia (excluding Papua Province, Waigeo, Aru
[but Kai is included]), and thePhilippines. They belong to 137
families. Their taxonomy and nomenclature are reviewed. Theoriginal
descriptions of all 7047 recorded species-group names and 1980
genus-group nameshave been checked in the original works for
correct spelling, types, type locality and bibliographicreferences.
The bibliography includes about 4700 titles. Synonymies are given,
based onpublished information as well as unpublished
observations.
The names of 49 introduced species and 347 extralimital taxa
cited in the discussions havealso been checked. The original
descriptions of all species not present in the covered area
butcited as type species of genera have been checked for
availability, authorship, date and correctspelling. The
availability of some family-group names has been checked when there
was suspicionof possible nomenclatural problems.
Bibliographic notes include new informations on the dates of
publication of works by, amongothers, Bleeker, Bloch, Heckel and
Steindachner and discussion of authorship of names in
variousworks.
The main nomenclatural acts are listed below:– type species
designation for: Crayracion, Eleotris Scopoli, Eleotris Walbaum,
Encheliopus
Cloquet, Gymnorhinus, Oonidus, Pristipoma Cuvier, Sargus
Gronow;– type species fixation under Code art. 70.3.1 for:
Bdellorhynchus, Desmoprenes, Innoculus,
Paraprotosalanx;– type species fixation under Code art. 70.3.2
for: Centrurophis, Ovoides Duméril, Pseudoscarus,
Rabula, Waitea;– lectotype designation for: Alausa
argyrochloris, Atherina endrachtensis, Barbus gardonides,
Barbus lateristriga, Betta patoti, Betta rubra, Carcharhinus
commersonii, Clupea cyprinoides,Clupea gigantea, Clupea
thrissoides, Crossochilus benasi, Cyprinus clupeoides,
Cyprinuslamta, Engraulis rhinorhynchos, Esox alepidotus, Esox
argenteus Gmelin, Esox argenteusSchneider, Equula longispinis,
Gobiomoroides piso, Gobius niger, Gonorhynchus
bimaculatus,Hemiramphus buffonis, Hemiramphus brevirostris,
Hemiramphus georgii, Hemiramphusrusselli Valenciennes, Johnius
cataleus, Lobotes auctorum, Neostethus borneensis,Parosphromenus
parvulus Foersch & Korthaus, Pellona leschenaulti, Raja
edentula, Rajaguttata Shaw, Raja narinari, Rasbora trilineata,
Synaptura achira, Teuthis brevirostris, Teuthisjavus;
– neotype designation for: Engraulis dussumieri, Lutjanus
gymnocephalus, Ovoides fasciatus,Ovum commersoni, Platygaster
megalopterus, Scarus schlosseri, Sciaena jaculatrix,
Sciaenapentadactyla;
– declaration as nomina protecta: Albula Scopoli, Aplocheilus,
Clupea quadrimaculata, Cyprinusbola, Hemiramphus georgii,
Hippocampus Rafinesque-Schmaltz, Kuhlia, Lateolabrax,Mastacembelus
erythrotaenia, Oligolepis, Pelates, Phyllopteryx, Platycephalus
japonicus,Puntius proctozysron, Raja uarnak Gmelin, Selaroides,
Toxotes microlepis Günther;
– declaration as nomina oblita: Albula Osbeck, Barbus
carassioides, Centranodon japonicus,Clupea mauritiana, Conorynchus,
Cyprinus goha, Gobileptes, Hemiramphus brevirostris,Hemiramphus
russellii van Hasselt, Hippocampus Perry, Leptaspis, Mastacembelus
catenatus,Odontopsis, Percalabrax, Platerome, Platysoma, Pristipoma
Quoy & Gaimard, Rajaommescherit, Raja scherit, Raja schoukie,
Sphyraena japonica Bloch, Toxotes microlepisBlyth;
Family Monacanthidae ............................. 469Family
Tetraodontidae .............................. 469Family Molidae
........................................ 481
Appendices
................................................................
481Acknowledgements
................................................... 484
BibliographyBibliographic notes
.............................................. 485Literature cited
..................................................... 505
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Kottelat: Inland fishes of Southeast Asia
4
– first reviser action on correct spelling of Barbus platysoma,
Bathygobius variabilis,Boleophthalmus novaeguineae, Euchiloglanis
dorsoarcus, Gazza equulaeformis, Kurtus,Leiocassis longispinalis,
Neocorassius, Pareuchiloglanis namdeensis, Raja uarnak Walbaum;
– first reviser action on precedence of simultaneous publication
of original descriptions of:Parosphromenus parvulus, Puntius
roloffi;
– first reviser action on precedence of simultaneous synonyms:
Chanodichthys over Pseudoculter,Thrissina over Xenengraulis and
Scutengraulis, Xenengraulis over Scutengraulis, Apistuslongispinis
over A. bougainvillii, Barbus balleroides over B. hypsylonotus,
Carassioidesmacropterus over C. argentea, Cyprinus jogia over C.
sutiha, Cyprinus pausius over C. musiha,Eleotris ophicephalus over
E. madagascariensis, Gobius caninus over G. quadriporus,Macropodus
yeni over M. nigrocorpus, Mystus pahangensis over M. johorensis,
Placocheilusbibarbatus over P. imbarbatus, Raja mula over R.
tajara, Rohita vittata over R. rostellatus,Tetraodon caria over T.
gularis;
– Chaetodon macrolepidotus Linnaeus, 1758 (now in Heniochus) has
precedence over C.acuminatus Linnaeus, 1758 as ruled by ICZN, 1912
[Opinion 40]. This Opinion has beengenerally ignored.
– Oxygastri of Bleeker (1860c) is not available because it is a
descriptive term, and not based onthe genus name Oxygaster;
– new genera: Desmopuntius (type species: Barbus hexazona Weber
& de Beaufort, 1912),Oliotius (type species: Capoeta oligolepis
Bleeker, 1853), Puntigrus (type species: Barbuspartipentazona
Fowler, 1934), Striuntius (type species: Barbus lineatus Duncker,
1904), Pao(type species: Tetraodon leiurus Bleeker, 1850).
The main unsolved nomenclatural problems are:– the type species
of Acanthurus is Naso unicornis and an application to ICZN is
needed to
retain the name for species currently called Acanthurus;– the
status of Siganus and Teuthis awaits a ruling by ICZN.
KEY WORDS. — freshwater fish, brackish water, mangrove,
estuaries, taxonomy, nomenclature,Southeast Asia, Singapore,
Indonesia, Malaysia, Thailand, Cambodia, Laos, Vietnam,
Myanmar,Manipur, Borneo, Sumatra, Java, Sulawesi, Maluku, Palawan,
Sundaland, Indochina, Mekong,Red River, Chao Phraya, Salween,
Irrawaddy
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THE RAFFLES BULLETIN OF ZOOLOGY 2013
5
Nowadays it seems fashionable among many zoologists, botanists,
anatomistsand physiologists to slightly look down upon this kind of
systematic research.In our opinion quite unjustly. Though formerly
this kind of investigation may
have been overestimated, at the moment one should not relapse
into the oppositeerror. Most likely we are now more than ever in
urgent need of accurate
descriptions of species […].
P. Harting, 1878, Pieter Bleeker'’s obituary
human eyes. To these species I have not given the benefit ofthe
doubt. The now fashionable discussions about crypticspecies do not
change the situation: one nucleotide does notmake a species, be it
ever so cryptic. Further, in fishes, thecryptic species discovered
by molecular techniques that Ihave been told about have been
cryptic not because taxono-mists could not distinguish them, but
because no trained tax-onomist ever had an opportunity to examine
them.
Another limitation of the precautionary approach in South-east
Asian fishes is shown by the huge number of 'new' fishspecies that
have been described from Vietnam in recentyears. Their description
is of a quality that makes it simplyimpossible to even guess
whether or not they might be val-id. The identity and possible
distinctness of most will re-main in limbo as long as they are not
competently re-de-scribed or evaluated.
Similarly a number of families recognised in recent timesare not
recognised. Cladistic molecular phylogeny (whichuses principles and
mathematical algorithms that were calledphenetics 30 years ago, an
approach rejected by cladistics)has the great particularism of
creating fluctuating and tran-sient phylogenies. There is even a
case of co-authors pub-lishing contradictory phylogenies
simultaneously in twopapers (Mayden & Chen, 2010; Tang et al.,
2010; see Britz& Conway, 2011a–b). This shows that it is
imprudent toinstantly adopt the latest theory and that naming every
littletemporary lineage uncovered by molecular analysis has lit-tle
justification.
For this catalogue I have examined personally the
originaldescriptions of all the species and genera recorded in
theinland waters of Southeast Asia, all their synonyms (mak-ing a
total of about 7047 nominal species and about 1980nominal genera).
I also examined the original descriptionsof the type species of all
genera if they were not known inthe area, and those of the 347 taxa
cited in the Taxonomicand Nomenclatural Notes but not present in
area. All citednomenclatural acts were checked. The availability of
all non-fish names cited as senior synonyms of fish genera
waschecked. Synonyms based on fossil taxa and which havenever been
used for recent taxa are not included.
INTRODUCTION
The present catalogue aims to present the state of the art ofour
knowledge of the diversity of freshwater fishes of South-east Asia.
Work began in 1986 when I compiled a list of thefreshwater fishes
of the Indochinese region (Kottelat, 1989).The list expanded when I
worked on a book on the fishes ofwestern Indonesia (Kottelat et
al., 1993; Kottelat & Whit-ten, 1996). Initially it was
intended to include only the fresh-water species, but the work for
the Indonesian fish bookrequired the inclusion of all species that
had been recordedin inland waters, that is, including estuaries,
most mangroves,etc. In September 2013, the list includes 3107 valid
nativespecies, in 707 valid genera and 137 families. Only
namedspecies are included. I am aware of about 300 species to
benamed soon or already on museum shelves and a fair num-ber of
synonyms to re-validate. I expect an additional 500species still
awaiting discovery in the wild. In addition, thereare 49 introduced
and established species.
This catalogue is not the ultimate inventory of the fishes
ofSoutheast Asian inland waters. Many discoveries are stillahead of
us and a great amount of work remains to be donebefore we reach an
acceptable level of knowledge.
Taxonomy and systematics have two main goals. One is pri-marily
of academic interest: the study of the diversity ofliving organisms
and their phylogenetic relationships. Theother is of immediate
practical interest: inventories, surveys,documentation of
biodiversity, and the compilation of iden-tification tools. For the
proper management of natural re-sources, we need information on
numbers of species andtheir identification now, not sometime in the
distant future.If definitive conclusions are not possible with the
availabledata, then tentative decisions are needed. As for other
com-ponents of environmental management strategies, the
pre-cautionary approach should be the rule. In the present
con-text, in case of doubt on the distinctness of two species,
theprecautionary approach would be to retain them as
distinctawaiting (possible) further research.
This precautionary approach, however, has its limits.
Thedevelopment of molecular techniques has led some to rec-ognise
as 'species' populations distinguished only by a fewnucleotides;
complex statistics have been used to justify therecognition of
'species' otherwise not distinguishable by
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Kottelat: Inland fishes of Southeast Asia
6
About 6000 publications (in 22 languages) have been ex-amined
and about 4700 relevant ones are listed. All citedreferences have
been checked in the original publications.The references or the
details that I could not check (for ex-ample, a stolen plate in the
only copy that I could access ofan antique book) are marked in bold
face (3 out of 4700titles). A few references have been checked by
trusted col-leagues.
After completion of the catalogue, all names have been
cross-checked against other databases. Especially, all names
werecross-checked against Eschmeyer's (2013) Catalog of Fish-es
(CoF) in 2001. For each entry in CoF disagreeing withmy data, the
data were verified again in the original descrip-tions and
literature. In 2010-2011 all names were cross-checked one more
time, and in cases of disagreement theoriginal literature was
checked for a third time. Out of theabout 9785 checked names, the
data (spelling, author, date,types, type locality, etc.) for about
3440 differs from thosein CoF (35 %). Most of the differences are
minor and oflittle or no nomenclatural consequences (mainly related
withtype localities), but a significant number of differences
areserious. The problems are more frequent with the
pre-1860non-English literature (type series, type localities,
type-spe-cies fixations, dates). In these times of on-demand
biodi-versity informatics there are too many assumptions madeabout
the quality of the data and there seem to have beenfew or no
efforts to carefully evaluate the contents of suchlarge databases.
An analysis of the types of differences anderrors is in
preparation.
A significant number of nomenclatural problems were dis-covered
and the application of the International Code ofZoological
Nomenclature (hereunder Code) results in a num-ber of nomenclatural
changes. In a few cases of changesaffecting family-group names or
widely used names, requestshave been presented to the International
Commission onZoological Nomenclature (ICZN) to retain these names
intheir current usage (Scatophagidae, Ephippidae, Kottelat,2010b;
Siganidae, Kottelat, 2013b; Mystus, Kottelat & Ng,2007). In
other cases, I considered that the name changesare minor and I
simply applied the Code. At the genus level,I consider that changes
resulting from the application of theCode do not create more
problems than do changes result-ing from the normal increase of our
taxonomic knowledgeby the discovery of new taxa, new characters,
etc. I consid-er it appropriate to ask the ICZN to retain the
current usagein cases of potential confusion resulting from the
discoveryof overlooked type species designations. But I consider
itunjustified to ask for the suppression of names (usually se-nior
homonyms and synonyms); to me, the argument of sup-pressing names
for the sake of stability of nomenclature infavour of reputedly
'well-known' names does not hold in ageographic area where new
discoveries still abound andwhere the taxonomic system is still
very unstable. Underthat logic, dozens of names should be
suppressed and thiswould affect the stability of nomenclature by
making thepurpose of the Code irrelevant. Also, writing
applicationsfor all these minor cases would mean more applications
thanthe ICZN receives in a year.
Limitations. — One of the limitations of this catalogue isthat I
started it in 1986; the work spanned 24 years and fouroperating
systems and unavoidably this caused slight inter-nal
inconsistencies in formatting. It has been updated con-tinuously so
that the technical content is not affected by this'formatting
evolution'. Further, between 1992 and today, twodifferent editions
of the International Code of ZoologicalNomenclature (ICZN, 1985,
1999) have been in use, whichdiffer slightly. I have tried to
update all entries affected bythe changes but I may have missed
some.
Another limitation is that the core target of this work andmy
own experience is the 'real' freshwater fishes. My treat-ment of
these taxa is probably close to complete. But I amlikely to have
missed some records of estuarine species, orsome literature. When I
encountered nomenclatural prob-lems concerning freshwater fishes I
had no hesitation in tak-ing the necessary actions to clear the
problems. When it cameto the same situation with estuarine taxa, I
tried to solve theroutine problems but decided not to address the
more com-plex ones. However, I discuss these cases and their
possiblesolutions where pertinent. The number of marine taxa
withnomenclatural problems was unexpectedly high, and manywell and
long known genus and family names are involved.
A potential for small errors arose late in the preparation
ofthis catalogue. I had long tried to confirm or revise the
chro-nology of the many papers published by Pieter Bleeker andit is
only late that I obtained the data to establish the se-quence of
publication of some 270 papers he published dur-ing his stay in
Java (Kottelat, 2011a) and of the Atlas ich-thyologique (Kottelat,
2013c). A number of the names cre-ated by Bleeker appeared more or
less simultaneously indifferent papers and journals. A consequence
of this revisedchronology is that the now-established dates of
availabilityof many names differ from those commonly recognised,
andthis has changed the precedence of the different descriptionof a
few species, which now may have a different type se-ries, or of
some new genera, which now may have differentoriginally included
species, thus potentially invalidatingearlier type species
designations. I have tried to eliminatethis risk but I expect that
some details would have escaped me.
Geographic and habitat coverages. — The geographiccoverage
includes all inland water bodies of Southeast Asiabetween (and
including) the Kaladan, Irrawaddy and the RedRiver drainages, the
small coastal drainages between the RedRiver and Hainan (included),
the whole Indochinese Penin-sula, Andaman and Nicobar Islands,
Indonesia (excludingPapua Province, Waigeo and Aru [but Kai is
included]), andthe Philippines (Fig. 1).
All freshwater species are included. Introduced species
thatbecame established are listed (marked by asterisks, *),
butwithout complete synonymies; only species that have es-tablished
reproducing populations are listed. Species in-habiting the
estuaries, brackish lower stetches of rivers,mangroves, etc. are
also included. Species occasionally re-ported in freswaters are
recorded, although some of therecords or identifications need
critical reevaluations (which
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THE RAFFLES BULLETIN OF ZOOLOGY 2013
7
was beyond the goals of this work). I preferred to be
tooinclusive than too exclusive. For these species too, synony-mies
are complete and include the nominal species describedfrom
freshwater as well as the marine ones; similarly, thegeneric
synonymies applying to these genera are complete.Through a lack of
familiarity with some groups I may haveoverlooked some
synonyms.
Listed names. — All known names are listed,
includinginfrasubspecific names, which are given in their
originalform. The only names that are not included are those
sub-specific or infrasubspecific epithets typicus when they
aremerely intended to denote the nominotypical subspecies;such
names are usually not nomenclaturally available andshould not be
used.
Spellings. — The headings of all generic and specific ac-counts
have the correct spelling of all valid names. In thesynonymies,
however, all names are given with their origi-nal combination
(except that interpolated subgeneric namesare omitted) and with
their original spelling, including mis-spellings, capitalised
letters, and diacritic marks [ü, è, ñ, etc.].Capitalised letters
and diacritic marks are not permitted bythe Code (arts. 27, 28,
Glossary) and must be corrected. In-correct original spellings are
used only in the synonymies
but they have been corrected in all other circumstances,
es-pecially in the discussions under Nomenclatural notes.
Families. — Families are listed following the sequence inNelson
(2006), except within Cypriniformes, for whichI follow Šlechtová et
al. (2007) and my personal experience.When there is disagreement
between authors with regard tothe limits of families or higher
categories, I generally fol-lowed common practice, but have noted
alternatives.
With a few exceptions I have not searched the synonymiesof
family-group names. Note that a family-group name keepsits original
author and date even if used at different ranks.For example
Leuciscini Bonaparte, 1835 retains Bonaparte,1835 as author, even
if treated as subfamily Leuciscinae orfamily Leuciscidae.
Genera and species. — Genera are listed in alphabeticalsequence
within family. Species are listed in alphabeticalsequence within
genera.
Entries for genera include the valid name of the genus (inbold,
as a heading), the name of the genus with the spellingin the
original description, the author, the year of publica-tion, the
number of the page with principal information. This
Fig. 1. The geographic area covered by the catalogue.
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Kottelat: Inland fishes of Southeast Asia
8
is followed by information on possible subgeneric status inthe
original description, type species, mode of designation,information
on possible nomenclatural acts associated withthe name, and
grammatical gender. This information is pro-vided for all names
considered to be synonyms, in chrono-logical sequence.
Entries for species include the valid name of the species
(inbold, as the heading), the name of the species as spelt in
theoriginal description, the author, the year of publication,
thenumber of the page on which the actual description starts(or
where the elements necessary to make the name avail-able occur) and
the number of the main illustrations (ignor-ing those showing maps,
anatomical details, portrait of col-lector, etc.). This is followed
by a block in parentheses withinformation on type locality and
primary types, and infor-mation on possible nomenclatural acts
associated with thename. If the name is based largely or totally on
references tothe older literature, this information is listed first
in the pa-renthesed block). This information is provided for all
namesconsidered to be synonyms, in chronological sequence.
Additional information, if needed, is listed under
Nomen-clatural notes and Taxonomic notes. When names are citedunder
Notes, which are not mentioned elsewhere in the text,I usually (but
not always) added the same data for that nameat the end of the
paragraph, in brackets.
Transliteration of non-Latin alphabets. — Author names,place
names, and journal names in non-Latin alphabets, andin languages
using other notations, have been transcribed;titles of books and
papers have been translated. When a tran-scription is used in the
original work (e.g. in the text, in anabstract, in a table of
contents), the same spelling is usedhere. There are some
inconsistencies as it happens that tran-scriptions or translations
used in abstracts or tables of con-tents may be different from the
actual title of a paper. Fre-quently, transcription systems have
changed with time andno standardisation has been attempted here.
Older bibliog-raphies or indexes may have used earlier
transcription sys-tems and I consider that a standardised use could
actuallycomplicate bibliographic search, especially for those
notfamiliar with these languages.
Unfortunately, some accents and diacritic marks may
havedisappeared as standard western European keyboards andsoftware
do not support them. This especially applies in thecase of the
Vietnamese alphabet.
Type localities. — The type locality is the locality at whichthe
holotype, lectotype or neotype was collected. Althoughmentioned in
the Code, the type locality has no nomencla-tural role. Simply, it
is a convenient wording, it is shorter tosay type locality than
'the locality at which the primary name-bearing type was
collected', or to give the locality data infull.
In case there is no primary type but a series of syntypesfrom
different localities, the type locality is the sum of allthe
localities of the syntypes, and all their localities are list-
ed (separated by a slash [ / ] where clarity requires it).
Invery few cases (when the list of localities of syntypes isvery
extensive), I have merely given a general descriptionof the
localities. Localities are usually given with the origi-nal
spelling; this sometimes results in different spellingsbeing used
for the same locality under different headings;I have tried to
introduce some consistency, but only in caseswhere I was certain
that the different spellings were reallyreferring to the same
place, or when the different spellingswere used for the locality of
the very same specimen, orreferring to the very same bibliographic
source. Alternativespellings or modern equivalents are given in
square brack-ets, but this has not been systematically attempted.
Localitydescriptions have been translated into English when
possi-ble and/or justified; in some cases, words meaning
river,lake, etc. are part of the name in the original language
andthey have not been deleted in order to avoid ambiguitieswhen
using local maps [but the word river, lake, etc. hasbeen added].
Local names have been used, except for a fewwell known rivers and
lakes with a common English nameused in international literature
(e.g. Mekong, Irrawaddy,Salween, Ganges, Red River). For most
localities, when fea-sible I have tried to add information on
present politicalentities (country, province, state, etc.) and
river basin as anaid to the reader. For larger topographic features
that haveseveral different names, a single one has been
consistentlyused; this especially applies to those traversing
differentcountries (e.g. Mekong River); I usually retained the
nameeasiest to find for readers not familiar with local toponymyor,
when it exists, the English name used in internationalliterature
(e.g. Salween River and not Nu Jiang, Salawin,Thanlwin, Salouen, or
fGyl mo rNGul chu [a transcriptionfrom Tibetan language]; Irrawaddy
and not Ayeyarwady;Red River and not Song Hong, Yuan Jiang or
Fleuve Rouge).
As the work on the check-list spanned more than 24 years,it is
likely that some of the earlier entries might be in a
slightlydifferent format than the latest ones.
Infrasubspecific names and nomina nuda having no nomen-clatural
status, they do not have type specimens and there-fore do not have
a type locality and I thus list only a 'local-ity', when justified.
Localities are usually not indicated forinfrasubspecific names
based on aberrant specimens; theyare given only if the name has
been created for a particulargeographical form.
When a neotype has been designated, the type locality is
thelocality of the neotype. The original type locality [the
local-ity of the primary type mentioned in the original
descrip-tion], if different, is usually listed in square brackets
in or-der not to lose that information.
When a lectotype has been designated, the type locality isthe
locality of the lectotype. The original type locality [thesum of
the localities of all the syntypes mentioned in theoriginal
description], is not listed, unless justified.
Linnean species, pre-Linnean literature, unpublishedsources. —
The identity and synonymy of species named
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THE RAFFLES BULLETIN OF ZOOLOGY 2013
9
by Linnaeus [Linné] and other early authors present partic-ular
problems, since many of them named species not onthe basis material
they personally examined but by referenceto earlier literature
sources. For example, in his Systemanaturae, Linnaeus (1758) based
most species names on ear-lier accounts by himself and others.
These have been tracedwhen possible and the exact bibliographic
references given.These secondary references have been examined too;
veryoften this actually creates additional problems because
thesesecondary authors themselves refer to older publications,
etc.I have not always included such secondary references andhave
only rarely searched the tertiary and earlier sources.Additionally,
several of these earlier works exist in variouseditions and
Linnaeus' (and other's) bibliographic referencesare not detailed
enough to decide which editions were used.For example, I have had
the rare opportunity of examiningside by side different editions of
Gesner's Nomenclator aqua-tilium and Fischbuch but could not find
all texts to whichLinnaeus referred. As these books usually are
considered tobe antiquities or collector-items, interlibrary loans
or photo-copies are not possible. Examination and comparison of
thevarious editions would mean travelling to a number of li-braries
and investing a lot of time and money, beyond thelimits of the
present work. Although there is an obvious his-torical and academic
interest, the utility of the exercise is notobvious in the context
of biological research and usable out-puts. In such instances, I
merely list the reference as givenby the original author, updated
into current bibliographic sys-tem. References to unpublished data
are usually omitted un-less they are relevant for nomenclatural
purposes [for exam-ple, reference to an unpublished figure of a
type]. Type lo-calities listed are those given by the author of the
new name;but the actual type locality is that listed by the
author(s) onwhich the account is based. Where holotypes are extant,
or iflectotypes or neotypes have been designated, the locality
ofthese specimens of course becomes the type localities. Toidentify
the type specimens of nominal species described bythese earlier
authors, one has to follow about the same pro-cedure, that is, to
find the specimens on which the accountscited by (e.g.) Linnaeus
are based (for examples, see Wheel-er, 1958, 1985, 1991; Fernholm
& Wheeler, 1983; Kottelat,2003c; Kottelat & Persat, 2005;
Kottelat & Freyhof, 2009).Again, this is a tedious and time
consuming task; I did searchsome such cases when this was essential
for solving nomen-clatural problems, but did not search in detail
all these cases.
(Contrary to a common belief, names created by Linnaeusare not
sacred, however, since Systema naturae has beendesignated by later
taxonomists as the starting point of to-day's nomenclatural system,
the names he used are the firstavailable names for the concerned
taxa. By the simple logicof the principle of priority they will
remain, regardless ofhow usable or informative the descriptions
are—and theyusually are useless without recourse to other
sources).
Incertae sedis, genera inquirenda, species inquirendae,nomina
dubia. — Incertae sedis are valid family, generaand species of
uncertain taxonomic position. Genera incer-tae sedis are listed at
the beginning of the Order or Familyto which they belong. For
example, Pimelodus javus clear-
ly is a member of Siluriformes, but cannot be placed in
anyfamily; it is therefore listed under Siluriformes, before
thefamily accounts.
Species incertae sedis are listed at the beginning of the
fam-ily to which they belong. Sometimes they are placed in thegenus
in which my experience or that of knowledgeablecolleagues suggests
they may belong. Alternatively, for spe-cies placed in genera to
which there is a suspicion they donot belong, the generic name is
placed in single quotationmarks (e.g. 'Genus' species), sometimes
with a commentunder Taxonomic notes. The fate of a species incertae
sedisis to be placed in a genus.
A species inquirenda (plural: species inquirendae) is a spe-cies
of doubtful identity. Often they can be placed in a ge-nus but the
description and the known material do not allowa decision as to
whether or not the species is valid. Suchnames are listed
immediately under the heading of the ge-nus to which they belong.
Species inquirendae that cannotbe placed in any genus are listed
immediately under the head-ing of the family to which they belong.
A species inquiren-da may have great similarity to a valid species
but its iden-tity may remain open to doubt; these are listed in the
synon-ymy of that species, and are indicated by a question mark
infront of the name. Some species inquirendae are poorly de-scribed
but are nevertheless tentatively accepted as possi-bly valid, for
example because an illustration in the originaldescription suggests
they may be valid; awaiting confirma-tion or a usable description,
they are listed as 'normal' spe-cies but with a question mark. It
is noteworthy that a sub-stantial number of the taxa described from
Vietnam in thelast 15 years falls into the category species
inquirendae.
The fate of a species incertae sedis is that future studies
willshow to which genus or family they belong. The fate of aspecies
inquirenda is to be redescribed and either found tobe a valid
species or a synonym of some other species.
A species inquirenda should not be confused with a nomendubium.
A nomen dubium (plural: nomina dubia) is a nameof doubtful
application that is impossible to link with aknown species, or that
may apply to several species. Typi-cally, a nomen dubium would have
been described in the18th or 19th century, with a few laconic
sentences includ-ing no diagnostic characters usable today, or
based on a paint-ing or on an artificially prepared specimen
(examples in-clude species of Tetraodontidae based on deformed
driedspecimens brought to Europe by seamen in the 18th centu-ry; or
many species described from Chinese paintings in the19th century:
these paintings usually were not based on agiven specimen but often
were an artistic or idealised viewof the species, copied from
earlier classical paintings, orsometimes simply imaginary). The
fate of a nomen dubiumis not to remain so, but to become either a
valid name or asynonym after either taxonomic examination or
appropriatenomenclatural decisions.
A genus inquirendum (plural: genera inquirenda) is a ge-neric
name that can be placed in a family but whose de-
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Kottelat: Inland fishes of Southeast Asia
10
scription and associated species (usually species inquirendaeor
nomina dubia) do not allow a decision as to whether ornot it is
valid. Such names are listed immediately under theheadings of the
family to which they belong.
BASIC PRINCIPLES OF NOMENCLATURE
A sad reality is that a majority of the users of scientific
names,especially those in the geographic area covered by this
list,have not had the opportunity to study the rules of
nomen-clature. This now also applies to most researchers
complet-ing their studies in western countries. For this reason it
seemsnecessary to start with a long introduction explaining
basicprinciples of nomenclature, terminology, and how to
under-stand the data in the present list. More experienced
readerswill probably not need to read this section.
The most basic principle of nomenclature is that it dealswith
only the names of organisms not with the organismsthemselves. The
confusion between animals and their namesmars many taxonomic
discussions and is becoming increas-ingly common and damaging. It
is of concern that even theeditors of some scientific journals are
no longer able to makethis distinction, especially in fashionable
areas like molecu-lar systematics.
Nomenclature is about the correct formation and treatmentof
names and the objective application of the 'legal' criteriaof a
code, irrespective of taxonomic concepts or philosoph-ical
approaches. Taxonomy is about the scientific study oforganisms and
includes a level of subjective interpretationof observations that
may differ among scientists.
Code. — Here, the word Code refers to the InternationalCode of
Zoological Nomenclature. The current (4th) edi-tion was published
in 1999 and superseded the previouseditions with effect from 1
January 2000. The Code is pub-lished under the responsibility of
the International Com-mission of Zoological Nomenclature (ICZN,
www.iczn.org),a body of zoologists (29 as of 2010), independent of
politi-cal or national entities. Under exceptional circumstances
andfollowing a prescribed procedure, the ICZN has the powerto
suspend the application of any of the articles of the Code.These
decisions (called Opinions and Directions) are pub-lished in the
Bulletin of Zoological Nomenclature.
Nomenclatural acts. — A nomenclatural act is any of pub-lished
act that affects the nomenclatural status of a scientif-ic name.
This includes the creation of names, emendations,designation of
types, rulings of the ICZN, etc. Nomencla-tural acts are valid if
they satisfy the provisions of the Code;they are invalid and must
be rejected if they do not followthe Code. Treating a name as a
subjective synonym is a tax-onomic act, not a nomenclatural
act.
Original descriptions. — Original description within themeaning
of the Code (arts. 10–20) refers to the first use (cre-ation) of an
available name.
Available name. — An available name is a name that satis-fies
the criteria of the Code and may be used for a validspecies. The
main criteria is that a new name must be ac-companied by a
description and the designation of a name-bearing type (type
specimen(s) for new species, type spe-cies for new genera; see
below). An available name is notautomatically a valid name.
Since 2012, the Code allows the publication of new namesin
electronic-only publications if they fulfill a number ofconditions.
Among them, the work must have an ISSN orISBN number, be archived,
and be registered in Zoobank(www.zoobank.org).
Valid name. — A valid name is the correct name applied toa
species. To be valid, a name must first be available. But
anavailable name is not automatically valid (junior synonymsare
available names but invalid). A 'valid name' should notbe confused
with a 'valid species'.
A nominal species is any of the available names created fora
species, irrespective of its validity. If a valid species has
xsynonyms, the valid name and the x synonyms are x+1 nom-inal
species.
Species-group names, genus-group names, family-groupnames. — The
species-group includes all the names of taxaof the rank of species
and subspecies. The genus-group in-cludes all the names of taxa of
the ranks genus and subge-nus. The family group includes all the
names of taxa rankedabove the genus-group: superfamily, family,
subfamily,tribes, etc.
Spellings. — A basic principle of nomenclature is that
theoriginal spelling (the ones created by the author in the
orig-inal description) must be retained. There are a few
excep-tions and the Code is very precise about which spellingsmust
be corrected (incorrect original spelling). There areno spelling
that may be corrected, there are only spellingsthat must be
corrected or that must not be corrected. Themain categories of
corrections is that if the species name isa Latin adjective it must
agree in grammatical gender withthe gender of the genus name.
Incorrect original spellingsshould never be used. Corrections of
incorrect original spell-ings allowed by the Code are called
justified emendations.Any intentional (explained) corrections not
allowed by theCode is an unjustified emendation and should never be
used;unjustified emendations are available names with their
ownauthor and date, are objective synonyms of the emendednames, may
be homonyms and may be used as substitutenames. They are included
in the list but I may have over-looked some. Any unexplained change
or error is called anincorrect subsequent spelling. Incorrect
subsequent spell-ings are not mentioned in synonymies, except if
they haveused erroneously at least occasionally (example:
Noema-cheilus and Nemachilus as commonly used incorrect spell-ings
of Nemacheilus).
Date of publication. — While in everyday's language
thepublication date is more or less equivalent to the date of
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11
printing, in the context of nomenclature, the Code definesthat
the date of publication is the date at which a work couldfirst be
obtained (for sale or by free distribution). The dateof publication
of a work is important to determine prioritybetween two synonyms,
two homonyms or two nomencla-tural acts. The Code rules that the
earliest one has priorityover the youngest one (senior synonym vs.
junior synonym;senior homonym vs. junior homonym). Even a
difference ofone day is enough to give priority. The Code art. 23.9
al-lows exceptions (reversal of precedence), but only if
veryprecise conditions are met.
The year of publication is retained as printed on the
publi-cation. It may happen, however, that the work appeared at
adate different from that printed on the publication. If a
dif-ferent date is documented by reliable information, this
datemust be retained for nomenclatural purposes. I have not
at-tempted to check the effective publication date of all
citedworks; this would have been very time and effort-consum-ing. I
have invested time for such search only when it wasnecessary to
establish the precedence between two works,or if it is was
necessary to determine if a given work ap-peared before or after
some of the dates important to theCode, or if there was a suspicion
that a stated date is farfrom the actual date. Otherwise, I
consider that usage of thedate printed on the publication is more
important than theactual publication date (e.g. for retrieval on
library shelvesor interlibrary loans).
Problems relating to dates associated with taxa are
discussedunder Nomenclatural Notes. Those associated with
specificpublications are mentioned in the Literature Cited
section,under the respective titles. Some of the more complex
casesare discussed separately: see Bibliographic Notes.
If nomenclatural acts are available from a valid electronic-only
publication, the date of publication is the date of
firstdistribution. New names and nomenclatural acts first
madeavailable in electronic publications as 'accepted manuscript'or
'uncorrect proofs' are not available. Taxonomists shouldnot
circulate manuscripts or proofs because this is a serioussource of
future problems.
Priority, precedence. — There is a subtle difference be-tween
priority and precedence. Priority indicates seniority,that a work,
a name or a nomenclatural act was publishedbefore another one.
Precedence indicates that a name mustbe used instead of another,
either by application of the prin-ciple of priority, or because
precedence is reversed for ex-ceptions prescribed by the Code or by
rulings of the ICZN.
Authorship. — The names of animals are usually followedby the
citation of their 'author'. For example the name of thecarp is
Cyprinus carpio Linnaeus, 1758, in which Linnaeusis the author of
the name C. carpio and 1758 the date of theoriginal description.
The citation of the author and year ismerely a bibliographic
reference. It is in no case an indica-tion of ownership.
Unfortunately, many scientists forget orignore the purpose of the
mention of the author's name andthis has sometimes resulted in
unjustified emotional reac-
tions of authors when one of 'their' species is treated as
aninvalid synonym by others. Somehow, naming a new spe-cies is the
formulation of an hypothesis and there is no shameif an hypothesis
is found to be erroneous. Vexed ego maynever accept a synonymy.
The name of an author is written in parentheses when thename is
moved to another genus by subsequent authors. Forexample, Ompok
bimaculatus (Bloch, 1794) was originallydescribed as Silurus
bimaculatus by Bloch (1794).
For nomenclatural purposes, the Code (art. 50) defines
theauthor(s) of a work as the person(s) who first published aname
in a way that makes it available. In most cases theauthor of a name
is the person whose name appears as theauthor of the book or
article. In some cases of works bymore than one author, if one
author only is responsible forthe name, then that person is author
of the name and thename is then cited in the format Barbus
binotatus Valenci-ennes, in Cuvier & Valenciennes, 1842. I
personally thinkthat this kind of citation of authorship is
contrary to the pur-pose of mentioning authors only as a
bibliographic refer-ence but introduces a ownership aspect; in this
example itseems even less desirable since the authors of the work
hadthemselves decided that the work would appear under bothnames.
In my eyes it serves no nomenclatural purpose tosearch who did what
in such a work (of course I understandthe historical interest). But
the Code says so and even theICZN has ruled so on this precise
case.
If it is established that a person other than the person namedas
the author(s) of a work is alone responsible for both thename and
the conditions making the name available, thenthat person is the
author of the name (often called second-ary author). For example,
in Schneider (1801), the nameand the description of Mugil
cirrhostomus are from unpub-lished notes of Forster. Schneider had
no specimen and noother source to describe the species; therefore
Forster isauthor of the name, and the name is cited as M.
cirrhosto-mus Forster, in Schneider, 1801. [Forster's manuscript
waslater published by Lichtenstein (1844) and the two texts canbe
compared.]
If an external person is author of the description (for exam-ple
personal notes) and the named author of the work citeshim and
proposes a name for the taxon, then the named au-thor of the work
remains author of the name. For example,Schneider (1801) used some
of Forster's descriptions butchoose other names (for example to
avoid homonymy); inthis case, Forster is not responsible for both
the name andthe conditions making it available, and therefore
Schneideralone is the author.
If an external person merely suggested a name for a
speciesdescribed in a work, this does not make him the author ofthe
name. The author of the work is responsible for the con-ditions
making the name available (i.e., the description, des-ignation of
types, etc.) and therefore is the sole author. Forexample, when
Valenciennes (1846) described Cobitis fas-ciata he commented that
in their notes the collectors (Kuhl
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Kottelat: Inland fishes of Southeast Asia
12
and van Hasselt) had a drawing of this fish, which they
hadlabelled Naemacheilus fasciatus. Kuhl and van Hasselt
trag-ically died before they could publish the description.
Twen-ty-four years later, Valenciennes used the name for the
newspecies and wrote the description himself. This makes himalone
the author. The species must be cited as C. fasciataValenciennes,
in Cuvier & Valenciennes, 1846 and not C.fasciata Kuhl &
van Hasselt, in Cuvier & Valenciennes, 1846.
Posthumous works belong to these categories but can bemore
complex. Sometimes it is simply a matter of some-body publishing a
completed manuscript by a deceasedfriend or colleague; in such
case, the author of the manu-script is clearly the author of the
work and of the names.The Code art. 50.1.1 explicitly mentions
"satisfying the cri-teria of availability other than actual
publication". This meansthat if both the names and the descriptions
of the new taxaare entirely the work of a deceased person, then he
is theauthor. To organise the actual publication is excluded
fromthe conditions of authorship.
If the description of a given species has been modified bythe
editor, both original writer and editor might be co-au-thor of the
name. If the description is completely rewrittenby the editor,
based on his own observations, then he is theauthor of the
name.
Examples: Forsskål died during a travel around the Red Sea.His
notes were later assembled, organised and published byNiebuhr,
using the names in Forsskål's notes and withoutwork on the content
of the text. Forsskål is author of thenames. For bibliographic
(librarian) purposes, he is also treat-ed as author the work (see
also Bibliographic Notes).
The manuscript of Bloch's Systema ichthyologiae was to-tally
rewritten by Schneider, who also added numerous spe-cies, the index
etc. Schneider is author of the work, as indi-cated on title page,
but Bloch is author of some taxa.Schneider explicitly indicated the
species he described.
When he died, Spix had not written the text on the fishes
hecollected in Brazil. He had supervised the preparation ofmost
plates and had named the species on the plates. Agas-siz was hired
to write a text that could be distributed withthe plates. He wrote
the descriptions of all species. For somehe ignored the names on
the plates and created new names,and he is author of these names.
For the species names thatappear only in plates, created by Spix,
because the platesalone are among the conditions sufficient to make
a name
available, Spix is the author. And in the cases Agassiz
wrotedescriptions and used the names created by Spix on theplates,
they are coauthors of the name.
Type species. — Each genus-group name has a type spe-cies. The
type species of a genus name is the species whosename determines
the validity of a genus. If several speciesare placed in genus X,
with type species Xx and this genusis later divided into two
genera, the genus which includespecies Xx will continue to be genus
X while the other ge-nus will have another name. If two genera have
the sametype species, they are objective synonyms. Genus-groupnames
proposed after 1930 without the fixation of a typespecies are not
available (Code art. 13.3) [note that art. 13.3requires that the
fixation be "in the original publication [Art.68]" and that art. 68
includes, as "type species fixed in theoriginal publication", those
established by original designa-tion, by monotypy, by absolute
tautonymy and by Linneantautonymy].
Type genus. — Each family-group name has a type genus.The type
genus is the genus whose name has been used toform the name of the
family. For example Silurus is the nameof the genus used to form
the family group names Siluridae,Siluriformes, Silurinae, etc. For
nomenclature purposes thesethree words are a single name. Whatever
the rank within thefamily-group, these names retain the same author
and date.
Type specimens. — Each species-group name has a type.The type of
a species name is the specimen on which thename is based; the
phrase name-bearing type is more ap-propriate but, in order to
simplify texts, is not usually used.The type specimen is the type
of a name, not of a species. Itis therefore erroneous to understand
the type as a 'model'representation of a species or a specimen to
which all spec-imens must be identical to be called the same
species. Thetype concept is a nomenclatural standard and totally
inde-pendent of any taxonomic judgements or philosophical the-ory.
The type is only used to objectively define to whichspecies the
name must be applied. If the type specimen ofthe name Yus belongs
to species 1, then the name of species1 is Yus. If the type
specimens of the names Yus and Xusbelong to species 1, then the
names Yus and Xus are syn-onyms (and the senior one has
priority).
Only primary types (name bearing types) are listed here.Primary
types are holotypes, lectotypes, neotypes and syn-types. Other type
categories recognised by the Code areparatypes and paralectotypes
but have no nomenclaturalfunction. Other 'type' categories (e.g.
allotypes, topotypes,paratopotypes, paraneotypes) are not
recognised by theCode, should not be used and are ignored here.
Among them,allotype is sometime used to designate one of the
paratypesof a sex different from that of the holotype; topotype is
usedas a shortened way to say 'a specimen collected at the
local-ity where the primary type was collected'.
The holotype is the specimen that has been explicitly
desig-nated so (or by a similar wording) in the original
descrip-tion by the original author, or the only specimen
available
author author of and name then authorof work conditions itself
of name is
is making name created byavailable is
A A A AA A B AA B A AA B B B, in AA A and B B A & B, in AA A
and B A A & B, in A
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THE RAFFLES BULLETIN OF ZOOLOGY 2013
13
to the author, if there is clear evidence that the author
basedthe nominal species on a single specimen. There is only
oneholotype per species. In all cases where there is clear
evi-dence that the author based the nominal species on morethan one
specimen (including literature records) but did notdesignate a
holotype, then all these specimens are syntypes.When it is not
possible to determine from the original de-scription if a name is
based on one or several specimens,I usually use 'types' or
'holotype?'.
If the nominal species is based on a specimen explicitly
des-ignated as holotype and a number of additional specimensare
also explicitly designated as types, these are paratypes;allotypes
are thus paratypes. The sum of the holotype+para-types or the sum
of the syntypes is called the type series. Insome cases (especially
for species described by P. Bleeker),I indicate the size ranges of
the type series in square brack-ets since this can be an important
tool to recognise them(example: syntypes [12, 41–43 mm SL]).
In cases where there is no holotype but only a series of
syn-types, one of the syntypes may be designated as lectotype;
itthen has the same value as the holotype. The remaining syn-types
then become paralectotypes and lose their status asprimary types.
Lectotypes are designated when it is demon-strated or suspected
that the type series includes more thanone species; it allows the
name to be definitively fixed tothe nominal species to which the
lectotype belongs. Inci-dentally, the designation of a lectotype
also restricts the typelocality to the locality of the lectotype,
excluding the local-ities of the other syntypes. Paratypes and
paralectotypes arenot listed hereunder as they are not name-bearing
types.
If none of the specimens of the original type series remains,or
if the holotype or lectotype no longer exist (they have notbeen
preserved, are lost, or destroyed) and if the name can-not be
unambiguously linked to a valid species, then (andonly then) a
specimen can be designated as neotype thatwill have the same
function as the holotype. Incidentally,the designation of a neotype
also restricts the type localityto the locality of the neotype. All
designations of neotypesthat do not fully satisfy these and several
other conditionslaid down in the Code are invalid and must be
ignored.
A number of neotype designations are invalid because theneed for
a neotype is not stated or demonstrated. This re-quirement did not
exist in the 1985 Code (art. 75(b)) and animplicit justification
was enough. The requirement becameexplicit in the 1999 Code, with
an added clause (art. 75.3.1)requiring "a statement that [the
neotype] is designated withthe express purpose of clarifying the
taxonomic status orthe type locality of a nominal taxon".
Unfortunately the 1999Code is missing a clause explaining what
should be done ofneotype designation validly made before 2000 but
invalidunder the 1999 Code because of the absence of the
state-ment. Formally, they become invalid because art. 86.3
statesthat all former editions of the Code have no force.
This I interpret as an oversight of the editors of the 1999Code
and is unintentional, otherwise it could be the cause
of very serious instability in some groups, which would
betotally against the spirit of the Code. Many of the
pre-2000neotype designations do not have the statement required
bythe 1999 Code. I have retained as valid the neotypes
validlydesignated before 2000 under the 1985 Code.
After the original description, it may be necessary for
laterauthors to re-examine the primary type of a nominal speciesin
order to decide to which taxonomic species it applies, forexample
in cases when several similar species are later dis-covered and the
original description does not mention thecharacters now decisive to
determine to which of these spe-cies the name must be applied. It
is, however, not a necessi-ty to examine a primary type if the
original description pro-vides all the information needed for
identification. In fact,types may be fragile specimens, and they
should not be han-dled if not justified, and persons without
experience shouldnot be permitted to handle them. Primary types
must bedeposited in museums or other responsible institutions
andwith staff able to conserve them and make them accessibleto
later researchers. Even if there is political pressure in
somecountries to consider types as national property, types donot
belong to a country but to science and must be accessi-ble to
competent scientists irrespective of their nationality.Neotypes, by
definition, must be deposited in a recognisedinstitution (e.g.
museum).
A number of species described by earlier authors do not
haveknown types or they have been lost since the original
de-scription. This does not affect the availability of a name.For
example, a specimen described in the field and latereaten by an
author remains the type specimen. Or a speci-men used as model for
a figure remains the type specimen,even if it has not been
preserved.
When known, institutions in which primary types are de-posited
are listed, together with register number and, whenknown, the
number of specimens in square brackets (exam-ple: AAA 1234 [2], BBB
1233 [1]). When the primary typeswere deposited in a collection but
cannot presently be locat-ed, the institution is listed as they may
still be present (mis-identified, misplaced, uncatalogued), or as a
starting pointfor further search. The source for the catalogue
number isgiven when it is not the original description; besides,
manyof those listed in the original descriptions have also
beenchecked in published catalogues or in the institutions
them-selves. When there is a series of syntypes, I listed thoseI
could trace in the literature, but made no effort to trace thewhole
series; this would have been tedious, many of themhaving possibly
been used for exchanges between institu-tions, etc. NT indicates
that there is no (or apparently no)preserved type material, LU that
there was apparently pre-served type material but that its
whereabouts are not known.A question mark in front of the
abbreviation of an institu-tion indicates that the type(s) is
possibly there or that thetype status of the specimen is not
certain.
Institutional abbreviations used in the text are listed
below.For institutions for which no abbreviations have been usedin
literature, the abbreviations follow current use by work-
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Kottelat: Inland fishes of Southeast Asia
14
ers at these institutions (where possible) or Leviton et
al.(1985; Leviton & Gibbs, 1988) or Eschmeyer (2010). I didnot
automatically follow Leviton et al. and Eschmeyer as astandard
because for non-US collections the abbreviationsthey list are often
not those actually used by the institutionsthemselves. In case the
abbreviations used in these lists dif-fer from those used by
workers at these institutions, I retainthe second one (as long as
they make sense and do not rep-resent personal, bureaucratic or
chauvinistic idiosyncrasy).
Synonyms. — The word synonym is used with the meaningit has in
the Code, that is a new name applied to a speciesthat already had a
name. Names erroneously used for a spe-cies other than the one
originally described under that nameare misidentifications.
Misidentifications are not synonymsand are not included in this
catalogue. The Code rules thatin case an author thinks that two
names actually refer to asingle species (i.e., they are synonyms),
the name publishedfirst (senior synonym) is the valid name; the
name publishedlater (junior synonym) is invalid (cannot be used).
The jun-ior synonym nevertheless remain available; should a
laterauthor find that the type specimens of the two names actual-ly
refer to different species, the junior synonym might beused again
(if it satisfies conditions set by the Code).
If the two synonyms are based on the same specimen (i.e.,they
have the same primary type), they are objective syn-onyms and the
junior synonym is invalid. If the two namesare based on different
primary types that an author consid-ers as belonging to a single
species, they are subjective syn-onyms (they are subjective because
this is the taxonomicjudgement of an author and other authors may
disagree; inthe opposite case, objective synonymy is a purely
nomen-clatural issue, not depending on of taxonomic judgement).
Homonyms. — Two available names with identical spell-ings and
created independently for different taxa are calledhomonyms. The
Code rules that the name published first(senior homonym) is the
valid name; the name publishedlater (junior homonym) cannot be used
and must be replaced.Junior homonyms are permanently invalid,
unless satisfy-ing some precise conditions of the Code.
In the species-group, two homonyms created in the samegenus are
called primary homonym. Example: Barbus yun-nanensis Fowler, 1958
is a junior primary homonym of Bar-bus yunnanensis Regan, 1904.
Two species names originally established in different gen-era
but later combined with the same genus name are calledsecondary
homonyms. Example: Crayracion fluviatilis var.ocellata
Steindachner, 1870 was not a homonym of Tetra-odon ocellatus
Linnaeus, 1758 when established. Later (in1975), it was treated as
a valid species of Tetraodon and itsname became T. ocellatus
(Steindachner, 1870), a juniorsecondary homonym of T. ocellatus
Linnaeus, 1758.
Replacement names. — If a species name becomes invalidbecause it
is a junior secondary homonym, it must be re-placed. The name used
for replacement is called substitute
name. The substitute name is the next oldest synonym. Ifthere is
no available synonym, then a new replacement nameis established.
The new replacement name takes the sametype as the replaced name.
In the example above, Dekkers(1975) treated Crayracion ocellatus as
a valid species ofTetraodon and made it a junior homonym of
Tetraodon ocel-latus. He created the new replacement name T.
steindachnerito replace the junior homonym.
A junior secondary homonym rejected and replaced before1961 is
definitively invalid (there may be exceptions; Codeart. 59.3). But
a junior secondary homonym rejected after1960 but later considered
to be in a genus different from thesenior homonym becomes valid
again (Code art. 59.4). Inthe above example,when Crayracion
ocellatus Steindachner,1870 was treated as a valid species of
Tetraodon it had to bereplaced by Tetraodon steindachneri Dekker,
1975. ButT. steindachneri is now considered to be a valid species
ofDichotomyctere and the senior synonym must be reinstatedand the
valid name is now D. ocellatus (Steindachner, 1870),not D.
steindachneri.
Occasionally, some authors have replaced names becausethey
overlooked an already available name that should havebeen used as
substitute name, or because they did not likean existing name, or
because they found it inappropriate, orto follow the nomenclature
rules at the time, or under politi-cal pressure. These replacement
names are invalid and can-not be used. These names are called
unnecessary replace-ment names. In the above example, after the
creation of thenew replacement name Tetraodon steindachneri
Dekkers,1975 it was discovered that the misidentified T.
biocellatusTirant, 1885 in fact was also a junior synonym of T.
ocellatus(Steindachner, 1870). Therefore T. biocellatus became
thevalid substitute name for T. ocellatus (Steindachner, 1870)and
T. steindachneri became a junior synonym ofT. biocellatus. An
later, after moving the species to Dycho-tomyctere as mentioned
above, the replaced secondary jun-ior homonym T. ocellatus was
reinstated as D. ocellatus(Steindachner, 1870).
Infrasubspecific names. — Infrasubspecific names arenames
originally intended for categories below the subspe-cies level (for
example: varieties, natios). These names arenot recognised as valid
by the Code. They are neverthelesslisted here. For infrasubspecific
names, I have indicated thelocality stated by the original author,
but have not listedmaterial. Even if these have sometime been
called type lo-calities and types in the literature, as the names
are not avail-able for zoological nomenclature, these 'type
localities' and'types' have no nomenclatural status.
Infrasubspecific names may become validated by a subse-quent use
as subspecies or species name. Whenever I foundan infrasubspecific
name validated this way I noted it; ifsuch subsequen