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The fishes of the Batang Hari drainage, Sumatra, with description of six new species

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The fishes of the Batang Hari drainage, Sumatra, with description of six new species

Abstract and Figures

Fish surveys were conducted between 1994 and 2003 in the Batang Hari drainage, Sumatra. The fish fauna of the drainage now includes a total of 297 species of which 48 are new records (45 of them new records for Sumatra). Six new species are described in the families Cyprinidae (Crossocheilus obscurus, Osteochilus kerinciensis, Pectenocypris micromysticetus), Nemacheilidae (Nemacheilus papillos) and Cobitidae (Pangio atactos, P. bitaimac). Crossocheilus pseudobagroides, Diplocheilichthys, D. jentinkii, Osteochilus scapularis, O. vittatoides, Leptobarbus rubripinna and Ras-bora hosii are revalidated. Lectotypes are designated for Labeo oblongus and Rasbora hosii. The identity of Osteochilus enneaporos, Nemacheilus longipinnis and Monotrete leiurus are discussed. A brief overview of M. leiurus suggests that M. bergii and M. ocellaris are valid species.
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Verlag Dr. Friedrich Pfeil
ISSN 0936-9902
Ichthyological Exploration
of Freshwaters
Volume 20
Number 1
An international journal for fi eld-orientated ichthyology
Ichthyological Exploration of Freshwaters
An international journal for fi eld-orientated ichthyology
Volume 20 • Number 1 • March 2009
pages 1-96, 51 fi gs., 10 tabs.
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Ralf Britz, Department of Zoology, The Natural History Museum, London, United Kingdom
Sven O. Kullander, Naturhistoriska Riksmuseet, Stockholm, Sweden
Helen K. Larson, Museum and Art Gallery of the Northern Territory, Darwin, Australia
Lukas Rüber, Department of Zoology, The Natural History Museum, London, United Kingdom
Ivan Sazima, Museu de Zoologia, Unicamp, Campinas, Brazil
Paul H. Skelton, South African Institute for Aquatic Biodiversity, Grahamstown, South Africa
Heok Hui Tan, Raffl es Museum of Biodiversity Research, National University of Singapore, Singapore
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Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1, pp. 13-69, 44 figs., 6 tabs., March 2009
© 2009 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
The fishes of the Batang Hari drainage, Sumatra,
with description of six new species
Tan Heok Hui* and Maurice Kottelat*
,
**
Fish surveys were conducted between 1994 and 2003 in the Batang Hari drainage, Sumatra. The fish fauna of the
drainage now includes a total of 297 species of which 48 are new records (45 of them new records for Sumatra).
Six new species are described in the families Cyprinidae (Crossocheilus obscurus, Osteochilus kerinciensis, Pectenocypris
micromysticetus), Nemacheilidae (Nemacheilus papillos) and Cobitidae (Pangio atactos, P. bitaimac). Crossocheilus
pseudobagroides, Diplocheilichthys, D. jentinkii, Osteochilus scapularis, O. vittatoides, Leptobarbus rubripinna and Ras-
bora hosii are revalidated. Lectotypes are designated for Labeo oblongus and Rasbora hosii. The identity of Osteochilus
enneaporos, Nemacheilus longipinnis and Monotrete leiurus are discussed. A brief overview of M. leiurus suggests
that M. bergii and M. ocellaris are valid species.
* Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singa-
pore, Lower Kent Ridge Road, Singapore 119260, Republic of Singapore. E-mail: dbsthh@nus.edu.sg
** Route de la Baroche 12, Case Postale 57, 2952 Cornol, Switzerland. E-mail: mkottelat@dplanet.ch
Introduction
Although ichthyological exploration and research
had been carried out in Central Sumatra for over
a centruy, it has not been extensive. The first
notable collections of freshwater fish in Central
Sumatra were reported by Pieter Bleeker (1852a-e,
1853a-b, 1854a-b, 1855, 1858a-b, 1859a, 1860a-b).
Odoardo Beccari made a small collection of
fishes in western Sumatra in 1878 (Vinciguerra,
1879). Gustav Schneider obtained a collection of
freshwater fishes from Central Sumatra during
the period 1897-1899 (Schneider, 1906), which was
documented by Walter Volz (1903a-b, 1907), who
himself collected in Sumatra in 1900-1902 while
he was employed as geologist by the Royal Dutch
Company for the Exploitation of Petroleum in the
Dutch East Indies (Volz, 1909). Bruno Machan
(1930) wrote on a collection of fishes from western
Sumatra made by J. Schild in 1899. Alfred C.
Harrison and H. M. Hiller made a small collection
of freshwater fishes from western Sumatra in
1901, which was described by Henry W. Fowler
(1904). Max Weber and Lieven F. de Beaufort
(1912) reported on a large collection of fishes from
the Kampar and Indragiri drainages obtained by
Alfred Maass in 1907. The last known major col-
lections were made from Bagansiapiapi near the
mouth of the Rokan River by J. D. F. Hardenberg
in 1929 and 1933 (Hardenberg, 1931, 1934).
Comparatively little has been done on the
ichthyological fauna of the northern and southern
ends of Sumatra. In the 1890s, Elio Modigliani
made a small collection of freshwater fishes from
northern Sumatra, which was reported by Al-
berto Perugia (1893). A small collection of fishes
was made by H. C. Kellers in 1925 from south
and southwestern Sumatra (Fowler & Bean, 1927).
14
Tan & Kottelat: Fishes of Batang Hari
WEST
SUMATRA
Sungai Dareh
Muara Bungo
RIAU
JAMBI
SOUTH
SUMATRA
SUMATRA
South China Sea
Muara Tebo
Sungai Alai
Kuala Tungkal
Pijoan
Muara
Kompeh
Sabak
Berbak
Muara Bulian
Muara Tembesi
Rantau
Rantau
Panjang
Panjang
Rantau
Panjang
Jambi
Bangko
Sarolangun
Sungai Penuh
Indian
Ocean
N
0 100 km
B
a
t
a
n
g
H
a
r
i
T
e
b
o
B
a
t
a
n
g
H
a
r
i
T
a
b
i
r
Danau Gunung Tuju
Danau Kerinci
T
e
m
b
e
s
i
B
a
t
a
n
g
H
a
r
i
M
e
r
a
n
g
i
n
T
e
m
b
e
s
i
A
s
a
i
Fig. 1. The Batang Hari drainage, Sumatra. Dashed areas: swamps (drawing by Kelvin Lim).
Fig. 2. Batang Hari River in Jambi town, riverbank with view of Angso Duo market (July 1997).
15
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Ernst Ahl (1934) published on a small collection
of fishes from Lake Toba (in northern Sumatra)
collected by August Thienemann in 1933; Thie-
nemann himself (1957) listed the material he
collected at other localities during the same trip.
More recently, Soetikno Wirjoatmodjo reported
on the fish fauna of the Alas river in northern
Sumatra (Wirjoatmodjo, 1987). The present review
does not include isolated smaller papers that
record just a few species.
Kottelat (1995a: 422) stated that the knowledge
of Sumatran fish fauna is poor, having no pub-
lished surveys of a single large drainage. Since
1984, separately or together, we have had the
opportunity to obtain various collections from
central Sumatra. The results have been compiled
in the first authors thesis (Tan, 1999a) from which
the present paper is partly derived. We present
here a list of the species known from the Batang
Hari drainage, highlight some new distribution
record, comment on the taxonomy of some spe-
cies, describe taxa new to science and present
various information on biology. The elasmo-
Fig. 3. Batang Merangin, immediately downstream of
Danau Kerinci (June 1994).
/
Fig. 4. Danau Souak Padang, an oxbow lake connected to Sungai Pijoan (November 1996).
16
branchs have been reported by Tan & Lim (1998),
the Siluriformes by Tan & Ng (2000) and the
Anabantoidei and Channoidei by Tan & Ng
(2005).
Geography
Sumatra is the second largest island on the
Sunda Shelf, with an area of 475 000 km
2
, more
than 1700 km long but only 350 km at its widest
point. The western margin of the island (the
outer margin of the Shelf) is mountainous, dom-
inated by the Barisan range, and of volcanic ori-
gin consisting of mainly two folds of mountains
with a valley in-between. There are several large
volcanic lakes scattered in this valley, namely
(from North to South) Toba, Meninjau, Singkarak,
Kerinci and Ranau. Several large rivers flow in a
northeastern direction out of the mountains, in-
cluding the Rokan, Kampar, Indragiri, Batang
Hari and Musi. Rivers draining the lakes or pla-
teaus in the mountain range pass through stretch-
es of high gradient on the way to the coastal plain.
An example is Batang Merangin draining the
Kerinci plateau, whose course drops by 300 m in
a 15 km distance. Much of Sumatra is perhumid,
and the original cover was tropical rain forest and
swamp forest along the northeastern coast. The
northern tip of Sumatra has pronounced seasons
(Loeb, 1935; Whitten et al., 1987; Rainboth,
1996a).
The Sunda Shelf is an extended continental
shelf, which is currently exposed as a series of
large islands (Greater Sunda Islands: Java, Su-
matra, Borneo). Molengraaff & Weber (1921) first
noted that the entire shelf might have become
exposed during the Pleistocene glacial periods.
The extent of continental surface exposure has
varied greatly during the Pleistocene and the
shallow sea floor that connects the islands is actu-
ally a system of drowned river valleys (de Beau-
fort, 1951). The paths of the drowned rivers
during the most recent sea regression indicate
that drainages on modern islands actually were
part of a continuous network with parts of other
islands. The southern side of Borneo and the
northern side of Java were drained by the East
Sunda River during the late Pleistocene. The
southern tip of Sumatra shared a drainage with
some of the northern Javanese rivers at the east-
ern most end of the island through the Sunda
Straits that nowadays separates Sumatra and Java.
The rivers from the western side of Borneo, Cen-
tral Sumatra, and the western tip of the Malay
Peninsula formed the West Sunda River. In the
northern Straits of Malacca, rivers of northern
Sumatra and western Malay Peninsula took a
northwesterly course towards the Indian Ocean.
See Voris (2000) for maps of Pleistocene sea lev-
els.
The major drainages in Central Sumatra in-
clude the Musi, Batang Hari, Indragiri and Kam-
par. All these rivers drain into the South China
Sea. The Batang Hari (Fig. 1) is about 600 km from
the headwaters of its longest tributary to the
river mouth (Fig. 2). Its drainages covers most of
Jambi Province and the southern part of the
highlands of Sumatra Barat Province (Fig. 3). The
remainder of Jambi Province is drained by a few
smaller coastal drainages (which are included in
the present review). The eastern and central por-
tions of Jambi Province encompass much low-
lying areas, originally made up of mainly tropical
rainforests, oxbow lakes (Figs. 4-6), freshwater
swamp forests (Figs. 7-9) and peat swamp forests
(Figs. 10-14). The western portion of Jambi Prov-
ince is situated about 1000-3000 metres above sea
level, in the highlands of the Barisan range. The
highland volcanic lakes of Kerinci (710 m a.s.l.;
Fig. 15) and Gunung Tujuh (1950 m a.s.l.; Fig. 16)
are some of the headwaters of the Batang Hari.
The lake of Gunung Tujuh is the highest lake in
Southeast Asia (Whitten et al., 1987: 402; Clarke,
2001).
Material and methods
All measurements are taken from the left side
(whenever possible) with digital calipers. Method
for taking counts and measurement follow Kot-
telat (2001), except for Nemacheilus which follows
Kottelat (1984, 1990). Fin ray counts were obtained
under transmitted light using a binocular dissect-
ing microscope. The last two branched dorsal and
anal rays articulating on a single pterygiophore
are counted as 1 {. Notation for vertebrae counts
follows Roberts (1989). Numbers in parentheses
following a particular count are the numbers of
examined specimens with that count.
Unless otherwise noted, identifications follow
Tan & H. H. Ng (2000) and H. H. Ng & Tan (2000)
for Siluriformes, Tan & P. K. L. Ng (2005) for
Anabantoidei and Channoidei, Compagno &
Roberts (1982) and Last & Stevens (1994) for
Tan & Kottelat: Fishes of Batang Hari
17
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Table 1. Fishes observed or collected in Batang Hari basin. * new records for Batang Hari; ** new records for
Sumatra (new records includes all species first obtained during our surveys, some of which already recorded in
Kottelat (2008), Kottelat & Tan (2008), Kottelat et al. (2006), Lim & H. H. Ng (2008), H. H. Ng (1996, 2009), H. H.
Ng & Kottelat (1998), H. H. Ng & Lim (2006), H. H. Ng & Tan (2000), Tan (1998, 1999b), Tan & Kottelat (1998,
2008), Tan & Lim (1998), Tan & H. H. Ng (2000) and Tan & P. K. L. Ng (1996, 2005).
Family Carcharhinidae
Carcharhinus leucas (Müller & Henle, 1839)
Family Pristidae
Pristis microdon Latham, 1794
Family Dasyatidae
Himantura signifer Compagno & Roberts, 1982
Family Osteoglossidae
Scleropages formosus (Müller & Schlegel, 1844)
Family Notopteridae
Chitala borneensis (Bleeker, 1851)
Chitala lopis (Bleeker, 1851)
Notopterus notopterus (Pallas, 1769)
Family Megalopidae
Megalops cyprinoides (Broussonet, 1782)
Family Anguillidae
Anguilla bicolor McClelland, 1844
Family Muraenidae
Gymnothorax tile (Hamilton, 1822)
Family Clupeidae
Clupeichthys goniognathus Bleeker, 1855
Clupeichthys perakensis (Herre, 1936)
Clupeoides borneensis Bleeker, 1851
Family Engraulididae
Coilia lindmani Bleeker, 1858
Lycothrissa crocodilus (Bleeker, 1851)
Setipinna melanochir (Bleeker, 1849)
Family Sundasalangidae
**Sundasalanx platyrhynchus Siebert & Crimmen, in
Siebert, 1997
Family Cyprinidae
Albulichthys albuloides (Bleeker, 1855)
Amblyrhynchichthys truncatus (Bleeker, 1851)
Balantiocheilos melanopterus (Bleeker, 1851)
Barbichthys laevis (Valenciennes, in Cuvier & Valenci-
ennes, 1842)
Barbonymus gonionotus (Bleeker, 1850)
Barbonymus schwanenfeldii (Bleeker, 1854)
Boraras maculatus (Duncker, 1904)
Crossocheilus cobitis (Bleeker, 1854)
Crossocheilus gnathopogon Weber & de Beaufort, 1916
Crossocheilus langei Bleeker, 1860
Crossocheilus oblongus Kuhl & van Hasselt, in van Has-
selt, 1823
**Crossocheilus obscurus Tan & Kottelat, 2009
**Crossocheilus pseudobagroides Duncker, 1904
Cyclocheilichthys apogon (Valenciennes, in Cuvier &
Valenciennes, 1842)
Cyclocheilichthys enoplos (Bleeker, 1850)
Family Cyprinidae (continued)
Cyclocheilichthys heteronema (Bleeker, 1854)
Cyclocheilichthys repasson (Bleeker, 1853)
Diplocheilichthys pleurotaenia (Bleeker, 1855)
**Eirmotus furvus Tan & Kottelat, 2008
**Eirmotus isthmus Tan & Kottelat, 2008
Epalzeorhynchos kalopterus (Bleeker, 1851)
Hampala ampalong (Bleeker, 1852)
Hampala macrolepidota Kuhl & van Hasselt, in van Has-
selt, 1823
Hypsibarbus huguenini (Bleeker, 1853)
Labeo chrysophekadion (Bleeker, 1850)
Labeo erythropterus Valenciennes, in Cuvier & Valenci-
ennes, 1842
Labiobarbus fasciatus (Bleeker, 1853)
Labiobarbus leptocheila (Valenciennes, in Cuvier & Va-
lenciennes, 1842)
Labiobarbus ocellatus (Heckel, 1843)
Leptobarbus hoevenii (Bleeker, 1851)
**Lobocheilos ixocheilos Kottelat & Tan, 2008
Lobocheilos schwanenfeldii Bleeker, 1854
Luciosoma setigerum (Valenciennes, in Cuvier & Valen-
ciennes, 1842)
Luciosoma spilopleura Bleeker, 1855
Luciosoma trinema (Bleeker, 1852)
Macrochirichthys macrochirus (Valenciennes, in Cuvier
& Valenciennes, 1844)
Malayochela maassi (Weber & de Beaufort, 1912)
Mystacoleucus marginatus (Valenciennes, in Cuvier &
Valenciennes, 1842)
Neobarynotus microlepis (Bleeker, 1851)
Osteochilus bleekeri Kottelat, 2008
Osteochilus borneensis (Bleeker, 1857)
Osteochilus intermedius Weber & de Beaufort, 1916
**Osteochilus kerinciensis Tan & Kottelat, 2009
Osteochilus melanopleura (Bleeker, 1852)
Osteochilus microcephalus (Valenciennes, in Cuvier &
Valenciennes, 1842)
Osteochilus scapularis Fowler, 1939
Osteochilus schlegelii (Bleeker, 1851)
Osteochilus spilurus (Bleeker, 1851)
Osteochilus vittatus (Valenciennes, in Cuvier & Valen-
ciennes, 1842)
Osteochilus waandersii (Bleeker, 1852)
Oxygaster anomalura van Hasselt, 1823
**Paedocypris progenetica Kottelat, Britz, Tan & Witte,
2006
**Parachela cyanea Kottelat, 1995
Parachela hypophthalmus (Bleeker, 1860)
Parachela oxygastroides (Bleeker, 1852)
**Pectenocypris micromysticetus Tan & Kottelat, 2009
Puntioplites bulu (Bleeker, 1851)
18
Fig. 5. Danau Arang Arang, an oxbow lake near Muara Kompeh (July 1997).
Fig. 6. Danau Arang Arang, an oxbow lake near Muara Kompeh (high water; May 1994).
Tan & Kottelat: Fishes of Batang Hari
19
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 8. Danau Kamining, near Muara Bungo (May 1994). Type locality of Pangio atactos.
Fig. 7. Swamp forest at Pijoan (November 1996).
20
Table 1. (continued).
Family Cyprinidae (continued)
Puntioplites waandersi (Bleeker, 1859)
Puntius sp. Bertam
Puntius sp. Kerinci
Puntius gemellus Kottelat, 1996
Puntius hexazona (Weber & de Beaufort, 1912)
Puntius johorensis (Duncker, 1904)
Puntius lateristriga (Valenciennes, in Cuvier & Valenci-
ennes, 1842)
Puntius lineatus (Duncker, 1904)
Puntius tetrazona (Bleeker, 1855)
**Raiamas guttatus (Day, 1870)
Rasbora argyrotaenia (Bleeker, 1850)
**Rasbora brittani Axelrod, 1976
Rasbora cephalotaenia (Bleeker, 1852)
Rasbora dorsiocellata Duncker, 1904
Rasbora dusonensis (Bleeker, 1851)
Rasbora einthovenii (Bleeker, 1851)
Rasbora elegans Volz, 1903
Rasbora gracilis Kottelat, 1991
Rasbora jacobsoni Weber & de Beaufort, 1916
Rasbora kalbarensis Kottelat, 1991
Rasbora kalochroma (Bleeker, 1851)
Rasbora myersi Brittan, 1954
Rasbora pauciperforata Weber & de Beaufort, 1916
Rasbora paucisqualis Ahl, in Schreitmüller, 1935
**Rasbora subtilis Roberts, 1989
Rasbora sumatrana (Bleeker, 1852)
Rasbora trilineata Steindachner, 1870
Rasborichthys helfrichii (Bleeker, 1857)
Rohteichthys microlepis (Bleeker, 1851)
Schismatorhynchus heterorhynchos (Bleeker, 1854)
*Sundadanio axelrodi (Brittan, 1976)
Thynnichthys polylepis Bleeker, 1860
Thynnichthys thynnoides (Bleeker, 1852)
Tor tambroides (Bleeker, 1854)
Trigonostigma hengeli (Meinken, 1956)
Family Balitoridae
**Barbucca diabolica Roberts, 1989
Homaloptera ocellata van der Hoeven, 1830
Homalopteroides nebulosus (Alfred, 1969)
Homalopterula gymnogaster (Bleeker, 1853)
Neohomaloptera johorensis (Herre, 1944)
**Neohomaloptera sp.
Family Nemacheilidae
**Nemacheilus cf. kapuasensis Kottelat, 1984
**Nemacheilus papillos Tan & Kottelat, 2009
Nemacheilus longipinnis Ahl, 1922
Nemacheilus pfeifferae (Bleeker, 1853)
Nemacheilus selangoricus Duncker, 1904
Family Vaillantellidae
**Vaillantella euepipterus (Vaillant, 1902)
Vaillantella maassi Weber & de Beaufort, 1912
Family Cobitidae
Acantopsis dialuzona van Hasselt, 1823
**Kottelatlimia katik (Kottelat & Lim, 1992)
Kottelatlimia pristes (Roberts, 1989)
Lepidocephalichthys tomaculum Kottelat & Lim, 1992
**Pangio alcoides Kottelat & Lim, 1993
Pangio anguillaris (Vaillant, 1902)
**Pangio atactos Tan & Kottelat, 2009
**Pangio bitaimac Tan & Kottelat, 2009
Pangio cuneovirgata (Raut, 1957)
Pangio malayana (Tweedie, 1956)
Pangio oblonga (Valenciennes, in Cuvier & Valenciennes,
1846)
Pangio semicincta (Fraser-Brunner, 1940)
Pangio shelfordii (Popta, 1903)
Family Botiidae
Chromobotia macracanthus (Bleeker, 1852)
Syncrossus hymenophysa (Bleeker, 1852)
Syncrossus reversus (Roberts, 1989)
Family Bagridae
Bagrichthys hypselopterus (Bleeker, 1852)
Bagrichthys macracanthus (Bleeker, 1854)
Bagrichthys macropterus (Bleeker, 1854)
Bagroides melapterus Bleeker, 1851
Hemibagrus hoevenii (Bleeker, 1846)
Hemibagrus aff. nemurus (Valenciennes, in Cuvier &
Valenciennes, 1840)
**Hemibagrus velox Tan & Ng, 2000
Hemibagrus wyckii (Bleeker, 1858)
**Hyalobagrus flavus Ng & Kottelat, 1998
**Leiocassis hosii Regan, 1906
Leiocassis poecilopterus (Valenciennes, in Cuvier & Va-
lenciennes, 1840)
Mystus abbreviatus (Valenciennes, in Cuvier & Valenci-
ennes, 1840)
Mystus bimaculatus (Volz, 1904)
Mystus castaneus Ng, 2002
Mystus singaringan (Bleeker, 1846)
Mystus wolffii (Bleeker, 1851)
Nanobagrus fuscus (Popta, 1904)
**Nanobagrus stellatus Tan & Ng, 2000
*Nanobagrus torquatus Thomson, López, Hadiaty & Page,
2008
**Pseudomystus heokhuii Ng & Lim, 2008
Pseudomystus leiacanthus (Weber & de Beaufort, 1912)
Pseudomystus rugosus (Regan, 1913)
Pseudomystus stenomus (Valenciennes, in Cuvier &
Valenciennes, 1840)
Family Siluridae
Belodontichthys dinema (Bleeker, 1851)
Ceratoglanis scleronema (Bleeker, 1863)
Hemisilurus heterorhynchus (Bleeker, 1854)
Hemisilurus moolenburghi Weber & de Beaufort, 1913
Kryptopterus bicirrhis (Valenciennes, in Cuvier & Valen-
ciennes, 1840)
Tan & Kottelat: Fishes of Batang Hari
21
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Table 1. (continued).
Family Siluridae (continued)
Kryptopterus cryptopterus (Bleeker, 1851)
Kryptopterus eugeneiatus (Vaillant, 1893)
Kryptopterus limpok (Bleeker, 1852)
Kryptopterus macrocephalus (Bleeker, 1858)
Kryptopterus palembangensis (Bleeker, 1852)
Kryptopterus schilbeides (Bleeker, 1858)
**Ompok fumidus Tan & Ng, 1996
Ompok leiacanthus (Bleeker, 1853)
Ompok rhadinurus Ng, 2003
Phalacronotus apogon (Bleeker, 1851)
Silurichthys indragiriensis Volz, 1904
Wallago leerii Bleeker, 1851
Family Schilbeidae
Laides hexanema (Bleeker, 1852)
Pseudeutropius brachypopterus (Bleeker, 1858)
Pseudeutropius moolenburghae Weber & de Beaufort, 1913
Family Pangasiidae
Helicophagus typus Bleeker, 1858
Helicophagus waandersii Bleeker, 1858
Pangasius djambal Bleeker, 1846
Pangasius nasutus (Bleeker, 1863)
Pangasius polyuranodon Bleeker, 1852
Pseudolais micronemus (Bleeker, 1847)
Family Akysidae
Acrochordonichthys rugosus (Bleeker, 1847)
**Akysis fontaneus Ng, 2009
**Akysis heterurus Ng, 1996
Breitensteinia cessator Ng & Siebert, 1998
Parakysis grandis Ng & Lim, 1995
Pseudobagarius macronema (Bleeker, 1860)
Family Sisoridae
Bagarius yarrelli (Sykes, 1839)
Glyptothorax platypogon (Valenciennes, in Cuvier &
Valenciennes, 1840)
Glyptothorax aff. schmidti (Volz, 1904)
Family Chacidae
Chaca bankanensis Bleeker, 1852
Family Clariidae
Clarias batrachus (Linnaeus, 1758)
Clarias leiacanthus Bleeker, 1851
Clarias meladerma Bleeker, 1846
Clarias nieuhofii Valenciennes, in Cuvier and Valenci-
ennes, 1840
Clarias olivaceus Fowler, 1904
**Encheloclarias cf. kelioides Ng & Lim, 1993
**Encheloclarias velatus Ng & Tan, 2000
Family Ariidae
Arius maculatus (Thunberg, 1792)
Batrachocephalus mino (Hamilton, 1822)
Cephalocassis borneensis (Bleeker, 1851)
Cephalocassis melanochir (Bleeker, 1852)
Cryptarius truncatus (Valenciennes, in Cuvier & Valen-
ciennes, 1840)
Family Ariidae (continued)
Hemiarius stormii (Bleeker, 1858)
Nemapteryx nenga (Hamilton, 1822)
Netuma bilineata (Valenciennes, in Cuvier & Valenci-
ennes, 1840)
Osteogeneiosus militaris (Linnaeus, 1758)
Plicofollis argyropleuron (Valenciennes, in Cuvier &
Valenciennes, 1840)
Plicofollis dussumieri (Valenciennes, in Cuvier & Valen-
ciennes, 1840)
Plicofollis polystaphylodon (Bleeker, 1846)
Family Plotosidae
Plotosus canius Hamilton, 1822
Plotosus lineatus (Thunberg, 1787)
Family Hemiramphidae
Hemirhamphodon pogonognathus (Bleeker, 1853)
Family Belonidae
Xenentodon canciloides (Bleeker, 1854)
Family Aplocheilidae
Aplocheilus panchax (Hamilton, 1822)
Family Synbranchidae
Monopterus albus (Zuiew, 1793)
Family Syngnathidae
Doryichthys deokhatoides (Bleeker, 1853)
Doryichthys martensii (Peters, 1868)
Family Ambassidae
**Gymnochanda filamentosa Fraser-Brunner, 1955
Gymnochanda limi Kottelat, 1995
Paradoxodacna piratica Roberts, 1989
Parambassis apogonoides (Bleeker, 1851)
Parambassis macrolepis (Bleeker, 1857)
Parambassis wolffii (Bleeker, 1851)
Family Datnioididae
Datnioides microlepis Bleeker, 1854
Datnioides polota (Hamilton, 1822)
Family Sciaenidae
Boesemania microlepis (Bleeker, 1859)
Panna microdon (Bleeker, 1849)
Family Toxotidae
Toxotes jaculator (Pallas, in Schlosser, 1767)
Toxotes microlepis Günther, 1860
Family Nandidae
Nandus nebulosus (Gray, 1835)
Family Pristolepididae
Pristolepis fasciata (Bleeker, 1851)
Pristolepis grootii (Bleeker, 1852)
Family Polynemidae
Polydactylus macrophthalmus (Bleeker, 1859)
Polynemus dubius Bleeker, 1854
Polynemus multifilis Temminck & Schlegel, 1843
22
Fig. 9. Lower part (in freshwater swamp forest) of stream connecting Danau Rasau (a lake on a peat dome) to
Batang Hari (May 1994).
Fig. 10. Sungai Bakung, a stream in flooded peat swamp forest near Arang Arang (May 1994).
Tan & Kottelat: Fishes of Batang Hari
23
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 11. Sungai Bakung, a stream in flooded peat swamp forest near Arang Arang (May 1994).
Fig. 12. Danau Rasau peat swamp forest (May 1994).
/
elasmobranchs, and Kottelat et al. (1993) for the
other taxa. Authors of scientific names of species
of the Batang Hari drainage are listed in Table 1
and are not repeated in the text.
Unless otherwise stated, our material is de-
posited in ZRC and CMK, with holotypes and
part of the paratypes of the new species in MZB.
Abbreviations: BMNH, Natural History Museum,
London; CAS, California Academy of Sciences,
San Francisco; CMK, collection of the second
author; MNHN, Muséum National dHistoire
Naturelle, Paris; MZB, Museum Zoologicum
Bogoriense, Bogor; RMNH, Nationaal Natuurhis-
torisch Museum, Leiden; UMMZ, University of
Michigan Museum of Zoology, Ann Arbor;
USNM, National Museum of Natural History,
Smithsonian Institution, Washington; ZMA, In-
stituut voor Systematik and Populatiebiologie,
Amsterdam; and ZRC, Raffles Museum of Bio-
diversity Research, National University of Singa-
pore.
24
Results
The species observed or recorded on the Batang
Hari drainage and the small coastal drainages
immediately adjacent are listed in Table 1. A total
of 297 species are recorded, of which 48 were then
new records (45 were new records for Sumatra).
Several of the new records have already been
announced in earlier publications by us or associ-
ates: Kottelat (2008), Kottelat & Tan (2008), Kot-
telat et al. (2006), Lim & H. H. Ng (2008), H. H.
Ng (1996, 2009), H. H. Ng & Kottelat (1998), H.
H. Ng & Lim (2006), H. H. Ng & Tan (2000), Tan
(1998, 1999b), Tan & Kottelat (1998, 2008), Tan &
Lim (1998), Tan & H. H. Ng (2000) and Tan & P.
K. L. Ng (1996, 2005). The following are notes and
comments on selected species.
Family Osteoglossidae
Scleropages formosus
Remarks. Systematics of the genus Scleropages
follows Kottelat & Widjanarti (2005). Scleropages
formosus was sighted once in 1996 on Jambi fish
market, as a cut up posterior section, for sale as
Table 1. (continued).
Family Blenniidae
Phenablennius heyligeri (Bleeker, 1859)
Family Eleotrididae
Bunaka gyrinoides (Bleeker, 1853)
Butis humeralis (Valenciennes, in Cuvier & Valenciennes,
1837)
Oxyeleotris marmorata (Bleeker, 1852)
Oxyeleotris urophthalmoides (Bleeker, 1853)
Family Gobiidae
Brachygobius doriae (Günther, 1868)
**Eugnathogobius siamensis (Fowler, 1934)
**Eugnathogobius cf. festivus Larson, 2009
Glossogobius giuris (Hamilton, 1822)
Parapocryptes serperaster (Richardson, 1846)
Periophthalmodon septemradiatus (Hamilton, 1822)
Family Gobioididae
Caragobius urolepis (Bleeker, 1852)
Family Helostomatidae
Helostoma temminkii Cuvier, 1829
Family Anabantidae
Anabas testudineus (Bloch, 1792)
Family Osphronemidae
Belontia hasselti (Cuvier, in Cuvier & Valenciennes, 1831)
Betta coccina Vierke, 1979
**Betta cracens Tan & Ng, 2005
**Betta falx Tan & Kottelat, 1998
Betta pugnax (Cantor, 1849)
**Betta raja Tan & Ng, 2005
**Betta renata Tan, 1998
**Betta simorum Tan & Ng, 1996
**Luciocephalus aura Tan & Ng, 2005
Luciocephalus pulcher (Gray, 1830)
Osphronemus goramy La Cepède, 1801
*Parosphromenus cf. bintan Kottelat & Ng, 1998
Parosphromenus sumatranus Klausewitz, 1955
Sphaerichthys osphromenoides Canestrini, 1860
Trichopodus leerii Bleeker, 1852
Family Osphronemidae (continued)
Trichopodus trichopterus (Pallas, 1770)
Trichopsis vittata (Cuvier, in Cuvier & Valenciennes,
1831)
Family Channidae
Channa bankanensis (Bleeker, 1852)
Channa cyanospilos (Bleeker, 1853)
Channa gachua (Hamilton, 1822)
Channa lucius (Cuvier, in Cuvier & Valenciennes, 1831)
Channa marulioides (Bleeker, 1851)
Channa melasoma (Bleeker, 1851)
Channa micropeltes (Cuvier, in Cuvier & Valenciennes,
1831)
Channa pleurophthalmus (Bleeker, 1851)
Channa striata (Bloch, 1793)
Family Chaudhuriidae
**Bihunichthys monopteroides Kottelat & Lim, 1994
**Chendol keelini Kottelat & Lim, 1994
**Nagaichthys filipes Kottelat & Lim, in Kottelat, 1991
Family Mastacembelidae
Macrognathus circumcinctus (Hora, 1924)
Macrognathus maculatus (Cuvier, in Cuvier & Valenci-
ennes, 1832)
Macrognathus tapirus Kottelat & Widjanarti, 2005
Mastacembelus erythrotaenia Bleeker, 1850
Mastacembelus unicolor Cuvier, in Cuvier & Valenciennes,
1832
Family Cynoglossidae
Cynoglossus feldmanni (Bleeker, 1854)
Cynoglossus waandersii (Bleeker, 1854)
Family Soleidae
Achiroides melanorhijnchus (Bleeker, 1851)
Brachirus panoides (Bleeker, 1851)
Family Tetraodontidae
Auriglobus amabilis (Roberts, 1982)
Monotrete leiurus (Bleeker, 1851)
Monotrete palembangensis (Bleeker, 1852)
Tan & Kottelat: Fishes of Batang Hari
25
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
food fish. Juveniles are in the aquarium-fish trade
in season, usually towards last quarter of the year.
The Batang Hari and Musi drainages are inhab-
ited only by the green form, Riau by a gold form.
The species is rare or uncommon, due to over-
collecting and habitat destruction.
Family Anguillidae
Anguilla bicolor
Remarks. Species of Anguilla had not yet been
recorded from eastern Sumatra; they are re-
stricted to the western, ocean-facing slope of the
island. We obtained specimens from the Sungai
Puneh market in Kerinci; they could possibly have
come from Padang or nearby streams draining to
the Indian Ocean. They were sold alongside spe-
cies which are locally caught in Kerinci, and
therefore we tentatively retain the species as
present in the highlands of Jambi.
Family Muraenidae
Gymnothorax tile
Remarks. Normally in estuarine habitats, but
observed in freshwater in the lower Batang Hari
still under tidal influence.
Family Clupeidae
Clupeoides borneensis
Remarks. Fresh specimens (Fig. 17) have a yellow
caudal fin with black distal margin, somewhat
similar to the coloration of Rasbora myersi. Ob-
tained so far only from market in Jambi.
Family Sundasalangidae
Sundasalanx platyrhynchus
Remarks. Sundasalanx was placed in the family
Clupeidae by Siebert (1997). Britz & Kottelat (1999)
considered that there is no data to support this
conclusion and retained Sundasalangidae as a
valid family in Clupeiformes.
We have two specimens only and their iden-
tity cannot be definitively cleared without damag-
ing them as identification requires examination
of osteological characters. They seem identical or
closely related to S. platyrhynchus, originally de-
scribed from the Kapuas drainage, Borneo. How-
ever, Sieberts (1997) description of S. platyrhyn-
chus distinguishes it only minimally from S. mi-
crops Roberts, 1981. Siebert only mentions that
the eye of S. microps is much smaller (approx.
3 % of SL,vs. 4-6 in his Table 4) and its pharyngeal
dentition is much smaller. His Table 4 does not
really list differences in pharyngeal dentition
(pharyngobranchial 3 [as Ipb 3 in Table] tooth
plate small with 1-2 teeth in S. microps, vs.
small with 2-3 teeth in S. platyrhynchus; Ipb 4
with tooth plate small vs. large). Small and
large are not defined, but in the text, the large
pharyngobranchial 4 tooth plate is described as
small, about | size of pharyngobranchial 4; the
condition in S. microps is not described. The type
series of S. platyrhynchus had earlier been re-
ported as Sundasalanx sp. by Kottelat (1991) who
pointed to some differences with S. microps but
considered it premature to described this limited
sample as a distinct species.
Material. ZRC 43027, 1, 16.4 mm SL; Sumatra: Pantai
Anduri, Batang Hari. – CMK 11178, 1, 23.0 mm SL;
Sumatra: Batang Hari in Kampung Rantau Panjang.
Family Engraulididae
Coilia lindmani
(Fig. 18)
Remarks. This species is not commonly encoun-
tered in the local markets. It was originally de-
scribed from the Musi drainage in South Su-
matra (Kottelat et al., 1993; Fig. 18).
Setipinna melanochir
Remarks. The Batang Hari collections include
specimens with black pectoral fins as well as
specimens with hyaline fins, which can otherwise
not be distinguished.
Family Cyprinidae
Balantiocheilos melanopterus
Remarks. Balantiocheilos melanopterus seems ex-
tinct in the Batang Hari drainage, apparently as
a result of overfishing and habitat destruction.
This species was widely distributed in Malay
Peninsula, Sumatra and Borneo. It is also re-
26
Fig. 13. Air Hitam peat swamp, Berbak (June 1996).
Fig. 14. Degraded peat swamp forest at Sabak (November 1996).
Tan & Kottelat: Fishes of Batang Hari
27
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 15. Danau Kerinci (June 1996).
Fig. 16. Danau Gunung Tujuh (June 1996).
28
ported as very rare in the Kapuas (Kottelat &
Widjanarti, 2005) and the related B. ambusticauda
is extirpated in the Chao Phraya and of uncertain
status in the Mekong (Ng & Kottelat, 2007). It is
still present in the Musi (Sumatra) (pers. obs.).
We have seen material collected in Jambi in 1909
by Moolenburgh (ZMA 115.913).
Barbonymus gonionotus
Remarks. Cultivated and found either naturally
or introduced throughout Southeast Asia. Regu-
larly seen in markets in Jambi but we have not
seen wild specimens.
Crossocheilus obscurus, new species
(Figs. 19a-b)
Holotype. MZB 10706, 108.1 mm SL; Indonesia:
Sumatra: Sumatra Barat: market at Kiliran Jao, a
village along road from Sungai Dareh to Solok;
H. H. Tan & H. H. Ng, 13 Dec 2003.
Paratypes. ZRC 49879, 2; UMMZ 248781, 2; CMK
18915, 2; 76.6-138.8 mm SL; same data as holo-
type.
Diagnosis. Crossocheilus obscurus is distinguished
from its congeners in South East Asia by the fol-
lowing combination of characters: size up to at
least 142 mm SL; one pair of rostral barbels, no
maxillary barbels; midlateral stripe with edges
not sharply contrasted, slightly curved down-
ward, obscured in largest individuals, continued
on median caudal-fin rays, reaching posterior
margin; no black mark between anus and anal
fin; mouth wide (30-36 % HL).
Description. General appearance in Figures 19a-b;
morphometric data of holotype and 6 paratypes
in Table 2. Dorsal fin with 3 simple and 8 {
branched rays; dorsal-fin origin above lateral line
scale 9 (3), 10 (3) or 11 (1). Pectoral fin rounded to
slightly falcate, with 15 (5) or 16 (2) rays, reaching
halfway to pelvic-fin base. Pelvic fin triangular
to slightly falcate, with 9 rays, reaching about
3
/
5
of distance to anal-fin origin and slightly beyond
anus; axillary scale present. Anus separated from
anal-fin origin by 3 scales (4 in one specimen).
Anal fin with 2 simple and 5 { branched rays.
Caudal fin with 10+9 principal rays, 9+8 branched.
Caudal peduncle 1.50-1.68 times longer than deep.
31-32 + 1-2 = 32-33 scales along lateral line, 8 or 9
predorsal scales, { 4/1/5 { scales in transverse
line, { 3/1/3 { scales in transverse line on caudal
peduncle, 3 or 3 { scales between lateral line and
pelvic-fin origin.
One pair of rostral barbels, length about { to
} of eye diameter. Upper lip with 14-18 fimbriae
(18 in 5 specimens), entirely covered by small
papillae. Anterior edge of lower lip with a few
large papillae.
Coloration. Preserved: Dorsal half of body, and
top and sides of head brown, all scale pockets
blackish. Ventral half of body and ventral and
lower surface of head yellow to pale brownish,
scales of upper 2-3 rows with brown margins.
A darker midlateral stripe along lateral line, ex-
tending on scale rows above and below lateral
line row. Edges of stripe not sharply marked,
somewhat irregular. All fins with hyaline mem-
branes, rays usually margined by a few black
pigments, denser at level of branching points.
Two median caudal-fin rays black, adjacent mem-
branes dark grey, continuing midlateral stripe.
Distribution. Crossocheilus obscurus is presently
known only from the middle Batang Hari drain-
age.
Etymology. Obscurus, Latin, dark, allusion to the
dark general appearance compared to other South
East Asian species of the genus. An adjective.
Remarks. The Batang Hari samples include
material of five species of Crossocheilus. The most
common one is a small species (up to 52 mm SL)
commonly found in the lowlands, in murky wa-
ters, usually with muddy bottom, under logs,
floating houses, boats, etc. It has two pairs of
barbels; the midlateral stripe continuous from tip
of snout to base of caudal fin, with a conspicuous
small blotch at posterior extremity, faintly marked
on caudal fin; a faint mark between anus and
anal-fin origin in juveniles; and a narrow mouth.
We identify it as C. cobitis (see below) (Fig. 20).
The second species, which we identify as
C. langei (Fig. 21), reaches 75 mm SL, has two pairs
of barbels; a well contrasted colour pattern with
the midlateral stripe continuous from the tip of
the snout to the tip of the median caudal-fin rays,
sometimes with a blotch at the posterior extrem-
ity of the caudal peduncle; sometimes a con-
spicuous black blotch between the anus and the
anal-fin origin; and a narrow mouth.
Tan & Kottelat: Fishes of Batang Hari
29
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
The third species, which we identify as C. pseu-
dobagroides (Fig. 23), reaches up to 52 mm SL, has
a body shape very similar to that of C. cobitis and
also occurs in lowland streams. It has one or two
pairs of barbels, a conspicuous black spot between
the anus and the anal-fin origin, up to four spots
on the dorsal mid-line and a conspicuous black
stripe ending in a blotch in the upper half of the
lower lobe of the caudal fin, not reaching the
posterior extremity of the rays.
The remaining two species have been col-
lected in swift waters over rocky bottom in the
foothills and hilly areas. The fourth species, de-
scribed here as C. obscurus, reaches 140 mm SL,
has a single pair of barbels; a midlateral stripe
from the opercle to the tip of the median caudal-
fin rays, slightly curved downward; the edges of
the stripe not sharply contrasted, not linear, and
obscured in the largest individuals; no mark
between the anus and the anal-fin origin; and a
wide mouth.
Finally, we group in a fifth species a number
of individuals reaching up to 90 mm SL, with a
single pair of barbels; a midlateral stripe from the
extremity of the opercle to the tip of the median
caudal-fin rays, conspicuously marked, no con-
spicuous spot at caudal-fin base; no mark between
the anus and the anal-fin origin; a small to mod-
erate mouth; and a short snout. We tentatively
identify them as C. oblongus (Fig. 22), but we are
not yet certain whether or not they are really
conspecific with the Javanese populations of
C. oblongus. The limited number of specimens we
obtained on markets, their preservation state as
well as the non-availability of recent material of
topotypical populations made an accurate iden-
tification impossible at the present stage.
Identification of the new Batang Hari species
(C. obscurus) requires a prior clarification of the
identity of C. oblongus, C. cobitis and C. langei,
which have often been confused. The name C. ob-
longus was first used in a letter by van Hasselt
(1823) published by Temminck. The description
Table 2. Morphometric data of Crossocheilus obscurus (holotype and 6 paratypes, ZRC 49879, UMMZ 248781,
CMK 18915) and C. gnathopogon.
C. obscurus C. gnathopogon
holotype range mean lectotype
MZB 49879 ZMA 113745
Standard length (mm) 108.1 76.6-138.8 91.6
In percent of standard length
Total length 132.2 129.8-134.6 131.8
Head length 22.5 22.2-24.4 23.2 22.1
Predorsal length 46.3 45.0-49.3 47.6 45.1
Prepelvic length 52.0 51.7-53.8 52.6 49.1
Preanal length 76.5 75.3-78.4 77.2 71.7
Head depth 15.0 13.9-16.8 15.4 15.1
Body depth at dorsal-fin origin 23.8 21.7-25.6 23.3 22.7
Depth of caudal peduncle 11.0 10.8-12.1 11.4 12.3
Length of caudal peduncle 18.5 17.7-19.0 18.1 20.1
Snout length 10.9 10.9-13.2 11.7 7.5
Eye diameter 5.4 4.8-5.9 5.4 5.2
Interorbital width 10.6 9.8-11.5 10.7 9.2
Mouth width 7.0 6.8-8.5 7.7 6.6
Distance from anus to anal fin 5.1
Dorso-hypural distance 60.7
In percent of head length
Snout length 49 48-54 51 34
Eye diameter 24 22-25 23 24
Interorbital width 47 43-48 46 42
Mouth width 29 30-36 32 30
Ratio length/depth of caudal peduncle 1.68 1.50-1.68 1.55 1.63
30
Fig. 17. Clupeoides borneensis, ZRC 41953, about 65 mm SL, fresh; Sumatra: Batang Hari drainage (July 1997).
Fig. 18. Coilia lindmani, ZRC 44138, about 150 mm SL, fresh; Sumatra: Batang Hari drainage (Oct 1999).
Fig. 19. Crossocheilus obscurus; Sumatra: Batang Hari drainage, Kiliran Jao; a, holotype, MZB 10706, 108.1 mm SL;
b, one of the paratypes, about 120 mm SL, fresh (Dec 2003).
b
a
Tan & Kottelat: Fishes of Batang Hari
31
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 20. Crossocheilus cobitis, ZRC 42376, 41.4 mm SL; Sumatra: Batang Hari drainage.
Fig. 21. Crossocheilus langei, ZRC 44141, 75.1 mm SL; Sumatra: Batang Hari drainage.
Fig. 22. Crossocheilus oblongus, ZRC 49228, 70.1 mm SL; Sumatra: Batang Hari drainage (right side, reversed).
is laconic but suffices to make the name available,
with Kuhl & van Hasselt as authors (Kottelat,
1987). It is impossible to know from van Hasselts
text on how many specimens his account was
based. Alfred (1971: 101) assumed that specimen
RMNH 1755 is the holotype. In fact, RMNH has
several specimens collected by van Hasselt (see
Roberts, 1993: 15), which probably are all syntypes
and Alfreds listing of one of them as holotype
makes it a lectotype by inference of a holotype
(ICZN art. 74.6). Roberts (1993: 15) listed specimen
RMNH 2640 as probable holotype or syntype,
but did not provide explanations.
Labeo oblongus Valenciennes (in Cuvier &
Valenciennes, 1842: 357) is based on the same
RMNH 1755 specimen and on a figure sent by
van Hasselt with his letter to Temminck. It is not
possible to determine if the model of the figure
shows the same specimen or another of the RMNH
specimens and therefore both are syntypes. We
32
designate RMNH 1755 as lectotype of Labeo ob-
longus Valenciennes and this makes it an objective
junior synonym and junior homonym of Crosso-
cheilus oblongus Kuhl & van Hasselt.
Van Hasselts drawing is reproduced in Rob-
erts (1993: 55, fig. 2), but the identity of the figured
fish is irrelevant for nomenclatural purposes as
the identity of the name is fixed by the lectotype.
Unfortunately we have not been able to examine
the lectotype. Nevertheless, it is worth mention-
ing that the figure shows a slender fish with a
blueish colour, a stripe not extending on the
caudal fin, and a relatively narrow mouth (not
quantified). The figure does not show the barbels,
but Alfred (1971: 102) observed only one pair.
This combination of characters corresponds to
our C. oblongus, except that our material has a
faint stripe on the caudal fin.
Alfred (1971) compared material from the
Malay Peninsula with the lectotype of C. oblongus
and concluded that they are conspecific and that
C. siamensis is possibly conspecific too. We exam-
ined part of the material in ZRC identified as
C. oblongus by Alfred. While these specimens
generally agree with the above diagnosis of our
C. oblongus, we observe variation in the shape of
the caudal peduncle, the width of the mouth and
the head, and the shape and pigmentation of the
caudal fin. The differences are observed both
between our Batang Hari material and these
Malayan samples identified by Alfred, as well as
within both sets. The differences we observe
among the Malayan material are as follows:
ZRC 2294, 9 specimens not clearly identifiable
(specimens in poor condition): ? C. cobitis
(long snout, caudal spot, 2 pairs of barbels)
and ? C. oblongus (short snout, no caudal spot,
1 pair of barbels);
ZRC 2296, 1 specimen: C. oblongus (long snout,
conspicuous caudal spot, faint stripe on cau-
dal, 1 pair of barbels);
ZRC 1953, 2 specimens: C. oblongus (long snout,
stripe conspicuous but not extending on cau-
dal fin, no caudal spot, similar to drawing of
C. oblongus in Bleeker [1863-64: pl. 105 fig. 2]
and in van Hasselts drawing [Roberts, 1993],
1 pair of barbels).
Our observations on other C. oblongus from the
Sundaic area suggest that a number of species
have been confused under the names C. oblongus
and C. siamensis (see also Kottelat, 2000a: 39).
While it is premature to discuss further their
identity, we can conclude that Alfreds concept
of C. oblongus (including its lectotype) refers to
fishes mostly similar to our C. oblongus as diag-
nosed above. For the time being, we tentatively
retain the name C. oblongus for both our Batang
Hari and Malayan material. Future studies may
show that the Malayan populations are specifi-
cally distinct. Kottelat (2000a) considered Indo-
chinese and Malayan populations as conspecific
with C.
oblongus, but this might have to be revis-
ited, once fresh material is obtained from Java. In
any case, with their linear, contrasted midlateral
stripe with sharp edges all these populations are
immediately distinguished from C. obscurus,
which has a midlateral stripe with edges not
sharply contrasted, not linear, obscured in largest
individuals.
Bleeker (1854a: 523) described C. cobitis on the
basis of 40 specimens 32-44 mm TL from Su-
matra (Padang) and Java (Batavia). The Atlas
account (Bleeker, 1863-64: 40, pl. 105 fig. 2) men-
tions 70 specimens 32-67 mm TL, including two
additional localities in Java (Krawang, Surabaya)
and there is no way to know exactly what is the
locality of the figured specimen. The drawings in
the Atlas usually are natural size (e.g., the figure
of C. langei [pl. 105 fig. 1, about 79 mm TL] shows
the only specimen known to Bleeker, which he
mentions (p. 42) to be 81 mm TL). The drawing
of C. cobitis is 68 mm TL and apparently shows
the longest of Bleekers specimens; thus, the
figured specimen is not a syntype and is from
either Krawang [Karawang] or Surabaya.
Bleeker (1854a, 1863) explicitly stated that
C. cobitis seems to always remain of small size,
and is distinguished from C. langei by a shal-
lower head (erroneous, see below), shorter barbels
and a wide silvery mid-lateral band with a me-
dian blackish line. The shape of the head disagrees
with Bleekers comment as it is more pointed and
shallower in C. langei (see below). We have seen
a single specimen labelled syntype of C. cobitis
(BMNH 1866.5.2.54), but we cannot be certain
that it is really a syntype. Also, it cannot be ex-
cluded that the remaining syntypes include more
than one species and a lectotype might be needed,
but it would be premature to designate it here.
Well developed tubercles in most specimens
of C. cobitis and the presence of ripe eggs in a
female 43.8 mm SL (ZRC 38828) are evidence that
they are sexually mature. It (or closely related
species) is known from Java, Sumatra, Borneo,
Malay Peninsula and Chao Phraya and Mekong
drainages.
Tan & Kottelat: Fishes of Batang Hari
33
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Bleeker (1860c: 127) described C. langei based on
a single specimen from Palembang, Sumatra. He
diagnosed it from his C. cobitis by having longer
barbels (two pairs), broad dark midlateral stripe,
a deeper head (1 { times in HL, vs 1 | to 1 ;
which in fact is erroneous, as 1.5 times in HL
means 67 % HL and 1.33-1.4 times means 71-
75 %). The description further states that the stripe
passes through a large black blotch at caudal-fin
base [not that it terminates at the blotch; basi
pinnae caudalis in maculam magnam nigram
transiente, transiens is the present participle
of transeo, to go through), and that there is a
round black blotch between the anus and the
anal-fin origin. This species has not been re-
ported again until Alfred (1971: 100) recorded it
from Malay Peninsula.
We have examined part of the material Alfred
identified as C. langei (ZRC 1783, 2, 90.5-105.4 mm
SL; ZRC 1788, 1, 70.7 mm SL). The specimens are
in a poor state, most of the pigments have disap-
peared, and we cannot see the black blotch be-
tween the anus and the anal-fin origin. Alfred did
not report the diagnostic blotch on the caudal-fin
base and we did not observe any remnant of it in
the damaged fins. Although Alfred reports the
presence of maxillary barbels, we did not see any,
but the mouth of all is damaged. ZRC 1788 and
one of ZRC 1783 (105.9 mm SL) seem conspecific
with our C. oblongus. The second specimen from
ZRC 1783 (89.3 mm SL) possibly belongs to
C. langei or C. gnathopogon; confirmation of the
presence of one or the other species in the Malay
Peninsula requires fresh material.
Two specimens from the Batang Hari (ZRC
44141, 69.6 and 75.1 mm SL; Fig. 21) agree with
Bleekers descriptions and figure of C. langei.
A third specimen (ZRC 43016, 93.5 mm SL) gen-
erally agrees too, but has a much fainter stripe
(possibly an artefact from poor preservation
[market-purchased specimen]) and a single pair
of barbels.
Weber & de Beaufort (1916: 234) distinguished
C. langei from C. cobitis by having only 2 scale
rows (vs. 3) between lateral lin and pelvic-fin
origin, and 4 { (vs. 5 {) below lateral line (appar-
ently in front of pelvic-fin origin). Our specimens
of the two species have 3 { and 5 { [in front of
pelvic-fin origin], respectively.
We identify as C. pseudobagroides Duncker, 1904
a small-sized species (up to 52 mm SL) with a
body shape very similar to that of C. cobitis. It has
one or two pairs of barbels, a conspicuous black
spot between the anus and the anal-fin origin,
spots on the dorsal mid-line and a conspicuous
black stripe ending in a blotch in the upper half
of the lower lobe of the caudal fin, not reaching
the posterior extremity of the rays. This is appar-
ently the undetermined species of Roberts (1989:
32, fig. 16c). It is presently known only from the
southern Malay Peninsula (Johor), Sumatra (Ba-
tang Hari), Kalimantan Tengah (Kahayan drain-
age) and Kalimantan Barat (Kapuas drainage)
(pers. obs.).
Crossocheilus pseudobagroides (Fig. 23) was
originally described from Muar River near Tubing
Tinggi, Malaysia (Duncker, 1904: 176). Dunckers
description and figure report diagnostic charac-
ters observable in our material: the distinct stripe
extending to the end of the upper rays of the
lower lobe of the caudal fin, with a blotch in the
middle of the caudal-fin base; the spots on the
dorsal mid-line (in front of dorsal-fin origin, at
dorsal-fin origin, at base of last dorsal-fin rays,
and behind dorsal fin); the black spot between
the anus and the origin of the anal fin; four barbels;
and 38 total lateral line scales. The spots along
the dorsal midline maybe very poorly contrasted
in specimens from turbid water (e.g. the specimen
on Fig. 23 shows only the spots in front of and at
dorsal-fin origin, and even these are poorly
marked). Alfred (1971: 100) treated C. pseudoba-
groides as a junior synonym of C.
langei, but part
of the material he listed as C. langei (ZRC 1783,
1788) belongs to C. oblongus or is not identifiable.
Besides, C. pseudobagroides differs from C. co-
bitis in having a greater predorsal length (dorso-
hypural distance falling much in front of tip of
snout, vs. about at tip of snout); a more slender
caudal peduncle (depth 1.63-2.04 times in its
length, vs. 1.51-1.68; length 17.5-19.6 % SL, vs.
16.3-19.1); more lateral line scales (33-34 + 2-3, vs
30-32 + 2-3); the anus closer to the origin of the
anal fin (3, vs. 4 scales; behind tip of pelvic rays,
vs in front); and a bolder and broader midlateral
stripe. The development of barbels seems variable,
with some specimens having rudimentary or
apparently no (e.g., ZRC 51770) maxillary barbels.
Crossocheilus cobitis also often has a few black
pigments between the anus and anal-fin origin,
but never forming a conspicuous blotch as in
C. pseudobagroides.
The last of the named Crossocheilus species from
Sumatra is C. gnathopogon Weber & de Beaufort,
34
Fig. 23. Crossocheilus pseudobagroides, ZRC 42748, 39.7 mm SL; Malaysia: Johor: Sungai Kahang.
Fig. 24. Crossocheilus gnathopogon, ZMA 113745, lectotype, 91.6 mm SL; Sumatra: Bukittinggi.
1916 (Fig. 24). It was described on the basis of two
specimens from Fort de Kock [Bukittinggi, Su-
matra]. Banarescu (1986: 150) also recorded spec-
imens from Bengkulu and Lake Kerinci. He re-
corded the type series as from Lake Mannindjan,
Fort de Kock [Lake Maninjau, Bukittinggi]. The
reason he mentioned Lake Maninjau is not known
and this mention is erroneous. There is neither
catalogue nor label data mentioning Lake Man-
injau (R. Vonk, pers. comm., 14 Nov 2008). It is
possible that Banarescu mentioned Lake Manin-
jau as the water body closest to Bukittinggi on a
large scale map. Waters from both Bukittinggi
and Lake Maninjau flow towards the southwest
to the Indian Ocean. Bengkulu is on the south-
western coast of Sumatra. The record from Lake
Kerinci requires confirmation.
Crossocheilus gnathopogon was diagnosed
Tan & Kottelat: Fishes of Batang Hari
35
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
mainly by the absence of rostral barbels. Weber
& de Beaufort reported the absence of rostral
barbel and the presence of minute maxillary
barbels. The lectotype (ZMA 113745; Fig. 24) in
fact has a tiny rostral barbel on each side in the
groove between the rostral cap and the posterior
part of the lower lip, and a small maxillary barbel
on the left side and none on the right side. This
is the specimen (with the dissected mouth) figured
by Weber & de Beaufort (1916: 232, fig. 96). The
paralectotype is figured by Kottelat et al. (1993:
pl. 7).
Fig. 25. Diplocheilichthys pleurotaenia; Sumatra: Batang Hari drainage, Sungai Dareh; a, ZRC 43003, 192.5 mm SL;
b, ZRC 51767, 108.0 mm SL, fresh (December 2003).
b
a
Fig. 26. Diplocheilichthys jentinkii; CMK 11525, 108.1 mm SL; Borneo: Kapuas drainage: Sungai Hulu Leboyan.
36
Key to the species of Crossocheilus
from Sumatra
1. - Up to 4 spots on dorsal mid-line; a black
spot between anus and anal-fin origin;
midlateral stripe ending in a bold blotch in
upper half of lower lobe of caudal fin, not
reaching posterior extremity of rays.
...........................................C. pseudobagroides
- No spot on dorsal midline; black spot bet-
ween anus and anal-fin origin absent (except
in C. langei); midlateral stripe ending at
caudal-fin base or at tip of median caudal-
fin rays.
........................................................................2
2. - Midlateral stripe ending at caudal-fin base,
with a small blotch at posterior extremity,
well marked on caudal peduncle, faintly
marked on caudal fin.
...........................................................C. cobitis
- Midlateral stripe ending at tip of median
caudal-fin rays, sometimes with a blotch at
posterior extremity of caudal peduncle.
........................................................................3
3. - Two pairs of barbels; a black blotch between
anus and anal-fin origin.
........................................................... C. langei
- A single pair of barbels; no black blotch
between anus and anal-fin origin.
........................................................................4
4. - Edges of midlateral stripe not sharply con-
trasted, irregular, not linear, and obscured
in largest individuals.
.......................................................C. obscurus
- Midlateral stripe linear, with sharp edges,
contrasted at all sizes.
.......................................................C. oblongus
Comparison material. The list includes only part of
the material we identified.
Crossocheilus cobitis: BMNH 1866.5.2.54, 1 syntype,
40.0 mm SL; Sumatra: Padang; or Java: Batavia [Ja-
karta]. – ZRC 42376, 18, 21.0-41.4 mm SL; ZRC 38550,
1, 52.0 mm SL; Sumatra: Batang Hari drainage. – CMK
11455, 11; CMK 11476, 2; Sumatra: Musi drainage. – CMK
11882, 1; Sumatra: Indragiri drainage. – ZRC 38828, 3,
37.9-43.8 mm SL; CMK 10427, 61; CMK 11611, 29; Bor-
neo: Kapuas. – CMK 13234, 7; Laos: Nam Leuk.
Crossocheilus langei: ZRC 44141, 2, 69.6-75.1 mm SL;
ZRC 43016, 1, 93.5 mm SL; Sumatra: Jambi: Angso Duo
market. – CMK 11761, 1; CMK 11675, 7; Borneo: Kapuas
drainage.
Crossocheilus gnathopogon: ZMA 113745, lectotype,
91.6 mm SL; Sumatra: Fort de Kock [Bukittingi].
Crossocheilus oblongus: ZRC 49288, 1, 70.1 mm SL;
Sumatra: Jambi. – ZRC 41550, 2, 56.1-89.8 mm SL; Su-
matra: Sumatera Barat: Sungai Dareh. – ZRC 2294, 9;
Malaysia: Negri Sembilan: Sungai Jelai. – ZRC 2296, 1;
Malaysia: Pahang: Sungai Tahan. – ZRC 1953, 2; Ma-
laysia: Pahang: Ulu Telom. – CMK 10555, 10; Borneo:
Kapuas drainage. – CMK 11858, 21; Borneo: Barito
drainage.
Crossocheilus pseudobagroides: ZRC 51770, 2, 44.1-
52.0 mm SL; ZRC 42748, 1, 39.7 mm SL; ZRC 42804, 1,
35.5 mm SL; Malaysia: Johor: Sungai Kahang. – ZRC
21102, 4, 30.5-40.0 mm SL; Malaysia: Johor: Sungai Se-
langi. – ZRC 51772, 37, 35.3-48.8 mm SL; Sumatra:
Jambi.
Diplocheilichthys pleurotaenia
Remarks. This species is redescribed by Kar-
nasuta (1993: 37) as Osteochilus pleurotaenia. Kot-
telat & Lim (1995b) commented that it should be
placed in a separate genus, for which the name
Diplocheilichthys Bleeker, 1860 is available. But the
material that Kottelat & Lim (1995b) referred to
as D. pleurotaenia in fact belongs to a distinct spe-
cies, originally described as O. jentinkii Popta,
1906 and later considered as a synonym of D. pleu-
rotaenia by Karnasuta (1993). Our comparison of
fresh material of both nominal species reveals
that they are clearly distinct from each other and
from all species of Osteochilus and we place them
in Diplocheilichthys. Lobocheilos rohitoides Bleeker,
1857 is a synonym of D. pleurotaenia.
Diplocheilichthys has the general appearance
of a typical Osteochilus but differs in details of
mouth and lip organisation. The mouth is much
wider than in species of Osteochilus (about 80 %
of interorbital width, vs. about 50 %), the inter-
orbital width itself is relatively wider (almost
equal to head width, as a result of a lateral exten-
sion of the frontal). The snout is conspicuously
protruding (vs. blunt). The lateral part of the
lower lip is plicated as in Osteochilus, but it is
concealed in the interior of the mouth (vs. ex-
panded laterally and exposed). Similarly, the
plicated part of the lower lip is not exposed later-
ally; only a few sparsely set papillae are exposed
on the anterior margin. The ventral surface of the
lower lip is hard and musculous. The rostral cap
entirely covers the upper lip in D. pleurotaenia or
leaves only the margin of the lip exposed in
D. jentinkii (vs. the basis only is covered in Os-
teochilus); the rostral cap has a lateral lobe (vs.
none).
Tan & Kottelat: Fishes of Batang Hari
37
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Weber & de Beaufort (1916: 214) placed
D. pleurotaenia in Labeo. They (1912: 524, 1916: 214)
redescribed L. rohitoides on the basis of a specimen
from Sumatra; they also mention having seen a
syntype of D. jentinkii. We found no difference
between their description of the Sumatra speci-
men and our material of D. pleurotaenia. They
redescribed D. pleurotaenia on the basis of litera-
ture accounts (Bleeker, 1855: 267, syntypes, from
Lahat; Volz, 1904, material from upper Langkat)
and specimen(s) from the upper and middle
Kapuas. We do not wish to discuss the identity
of the material in the references they cite. But
from the published data and our material, D. pleu-
rotaenia is unambiguously recorded from Java
and Sumatra. We have examined two juvenile
specimens from the Kapuas (CMK 6448, 11680)
that we identify as D. pleurotaenia. Diplocheilichthys
jentinkii is known only from the Kapuas drainage
(Borneo). Record of D. jentinkii from the Mahakam
(Roberts, 1989: 52) results from the erroneous
placement of the type locality (Bongan). The two
species of Diplocheilichthys are distinguished as
follows:
D. pleurotaenia (Figs. 25a-b): in lateral view,
snout pointed, almost triangular; head and body
dorsal profile steeply slanted; in lateral view, eye
not reaching dorsal profile; numerous small tu-
bercles on tip of snout; mouth arched, lower lip
rectangular, length about { of width; rostral cap
covering median part of upper lip; tip of anal fin
reaching end of caudal peduncle; depth of caudal
peduncle 1.2-1.4 times in its length; pectoral fin
reaching about } of distance to pelvic-fin origin;
4 { scale rows between lateral line and pelvic-fin
origin; body depth 30.7-32.9 % SL. See also Ta-
ble 3.
D. jentinkii (Fig. 26): in lateral view, snout
deep, squarish; head and body dorsal profile
glently slanted; in lateral view, eye almost reach-
ing dorsal profile; no tubercle on snout; mouth
strongly arched, lower lip squarish, length about
} of width; rostral cap not covering upper lip;
tip of anal fin not reaching end of caudal pedun-
cle; depth of caudal peduncle 1.7-1.8 times in its
length; pectoral fin almost reaching pelvic-fin
origin; 3 { scale rows between lateral line and
pelvic-fin origin; body depth 22.9-24.9 % SL. See
also Table 3.
See also discussion under Osteochilus scapula-
ris.
Material. Diplocheilichthys jentinkii: ZRC 49225,
3, 58.3-164.2 mm SL; CMK 16906, 1, 143.4 mm SL;
Borneo: Kalimantan Barat: Sungai Sibau. – CMK
11525, 9, 87.3-137.7 mm SL; Borneo: Kalimantan
Barat: Sungai Hulu Leboyan.
Diplocheilichthys pleurotaenia: ZRC 42349, 2,
60.3-58.5 mm SL; Sumatra: Jambi: Batang Hari at
Perlingat. – ZRC 42371, 2, 50.4-65.6 mm SL; Su-
matra: Jambi: Batang Hari at Desa Lounderang.
– ZRC 43003, 1, 192.5 mm SL; Sumatra: Sumatera
Barat: Sungai Dareh. – ZRC 51767, 2, 99.3-
Table 3. Meristic and morphometric data of Diplochei-
lichthys pleurotaenia (ZRC 43003 [1], ZRC 51767 [2], ZRC
51768 [1]) and D. jentinkii (ZRC 49225 [2]).
D. pleuro- D. jentinkii
taenia
Standard length (mm) 99.3-192.5 121.0-164.2
Meristics
Dorsal-fin rays iii, 11 { iii, 11 {
Caudal-fin rays 10+9 10+9
Anal-fin rays iii, 5 { iii, 5 {
Pelvic-fin rays 9 9
Pectoral-fin rays 16 15
Lateral line scales 30 + 2-3 30-31 + 3
Scale rows between dorsal-fin
origin and lateral line { 6 { 6
Scale rows around caudal
peduncle 15-16 (16) 15
Morphometrics
In percent of standard length
Total length 137.3-147.3 140.3-142.6
Head length 24.0-27.5 25.6
Predorsal length 45.7-48.7 43.4-44.6
Prepelvic length 51.5-53.8 49.9-50.5
Preanal length 74.4-78.8 74.8-75.4
Body depth at dorsal-fin origin 30.7-32.9 22.9-24.9
Body depth at anus 23.1-24.3 15.6-17.4
Depth of caudal peduncle 12.9-14.5 10.6-11.0
Length of caudal peduncle 15.7-17.2 17.7-18.3
Length of dorsal-fin base 20.8-25.0 20.4-22.7
Length of anal-fin base 9.4-10.8 9.0-9.2
Length of pectoral fin 22.3-26.4 22.7
Length of pelvic fin 23.3-27.0 22.8-24.6
Length of upper caudal lobe 39.8-45.4 43.2-43.3
Length of lower caudal lobe 36.6-45.1 42.4-44.0
Snout length 12.6-14.0 12.4-13.5
Eye diameter 5.0-6.7 4.9-6.4
Postorbital length 6.5-8.2 5.9-7.0
In percent of head length
Snout length 48-53 48-53
Eye diameter 21-26 19-25
Postorbital length 25-30 19-27
38
Fig. 27. Labeo erythropterus, about 800 mm SL, 7 kg, not preserved (December 2003); Sumatra: Batang Hari drain-
age, Pulau Punjung market (upper photograph by Ng Heok Hee).
a
b
108.0 mm SL; West Sumatra: Kiliran Jao. – ZRC
51768, 1, 105.1 mm SL; West Sumatra: Pulau
Punjung. – CMK 6848, 1, 41.1 mm SL; Borneo:
Kalimantan Barat: Kapuas River mainstream at
Nanga Embaluh. – CMK 11680, 1, 50.8 mm SL;
Borneo: Kalimantan Barat: Sungai Lanjak and
Sungai Lanjak Deras.
Labeo erythropterus
Remarks. Recorded from Air Penatai in Kerinci
(ZMA 120.816, 395 mm) by Roberts (1993). We
have examined one fresh specimen (about 800 mm
SL, 7 kg) at Pulau Punjung market at Sungai Dareh
(Fig. 27), which fits Bleekers (1863-64: 52, pl. 106)
description and plate.
Tan & Kottelat: Fishes of Batang Hari
39
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
a
b
Fig. 28. Leptobarbus hoevenii; a, ZRC 49880, 33.0 mm SL; Sumatra: Batang Hari drainage, Jambi; b, CMK 11721,
137 mm SL; Borneo: Kapuas drainage, Danau Sentarum.
Fig. 29. Leptobarbus rubripinna, ZRC 49881, 135.4 mm SL; Thailand: aquarium trade.
Leptobarbus hoevenii
Remarks. Fry of Leptobarbus hoevenii is harvested
from the wild and exported for growing out in
aquaculture. It is often encountered in the markets
as large uniformly sized individuals. The juveniles
from Sumatra look very different from those from
Indochina, in having a stouter body and smaller
eye.
Leptobarbus hoevenii is reported from Java,
Sumatra, Borneo, the Malay Peninsula, and the
Mekong and Chao Phraya drainages. Bleeker
(1851a: 207, 1863-64: 116, pl. 132 fig. 2) and Weber
& de Beaufort (1916: 96) described and figured
the species (on the basis of Sundaic material) as
plain coloured, with a black blotch behind the
opercle. Our material from the Batang Hari as
well as from Borneo and the Malay Peninsula
40
(ZRC 2303, 10, collected in 1968, before the begin-
ning of the importation of large quantities of fry
from Thailand) agrees with their descriptions.
Specimens 30-60 mm SL (ZRC 49880, 41947) have
a narrow faint stripe running along the flank,
between the lateral line scale row and the row
above it. This stripe is missing in specimens over
about 100 mm SL (ZRC 38849, 38987, 39040, 39150,
43163, CMK 4761, 11883, 11721) (Figs. 28a-b).
The material from Thailand (Mae Khlong,
Chao Phraya and Mekong drainages) is immedi-
ately distinguished from the Sundaic one by the
presence of a broad midlateral stripe covering
most of the scale row above the lateral line scales
(Fig. 29). The two species also differ in head shape.
In specimens up to at least 160 mm SL, the inter-
orbital area is clearly rounded in the Indochinese
specimens (vs. almost flat in the Sundaic ones),
in lateral view the head dorsal profile is slightly
rounded and symmetric to the ventral profile (vs.
straight to slightly concave), the eye is located at
mid-depth of head (vs. closer to the dorsal profile),
the barbels shorter (posterior maxillary one reach-
ing vertical through middle of eye, vs. beyond
posterior margin of eye), the pelvic and anal fins
are orange to red (vs. dark grey). The rounded
caudal lobes and anal fin of the figured specimen
from Thailand (Fig. 29) is possibly due to some
extent to it having been in captivity for some time;
nevertheless, Smiths (1945) figure shows them
more rounded than in the Sundaic material.
We consider that they are distinct species. The
Sundaic one is L. hoevenii whose type locality is
Banjarmasin (Borneo). The Indochinese species
is L. rubripinna (Fowler, 1937) whose type local-
ity is Kemarat (Thailand, Mekong drainage). The
photograph of L. hoevenii in Kottelat et al. (1993:
pl. 10) shows a specimen of L. rubripinna from
Thailand and the one in Kottelat (2001: 60; here
Fig. 28b) a specimen of L. hoevenii from the Kapuas
drainage (Borneo).
Leptobarbus rubripinna is cultivated in Thailand
and is exported to Malaysia. The fry is also report-
edly harvested in the wild in Laos for export. In
Sumatra, the fry of L. hoevenii is harvested for
export to Malaysia.
Material. Leptobarbus hoevenii: ZRC 38987, 2, 126.2-
149.1 mm SL; ZRC 43163, 3, 170-215 mm SL; ZRC 49880,
27, 28.0-34.3 mm SL; ZRC 50849, 2, 135.2-146.1 mm SL;
Sumatra: Jambi. – CMK 4761, 3, 107.9-133.1 mm SL;
Sumatra: Jambi. – ZRC 39150, 1, 104.9 mm SL; Sumatra:
Berbak. – ZRC 41947, 10, 40.1-63.8 mm SL; Sumatra:
Batang Hari in Kampung Rantau Panjang. – ZRC 39040,
2, 113.8-121.2 mm SL; CMK 11883, 1, 97.9 mm SL; Su-
matra: Riau: Indragiri drainage: Sungai Bengkwan. –
ZRC 2303, 10, 108.8-162.0 mm SL; Malaysia: Pahang:
Tasek Bera. – CMK 11721, 2, 137.0-145.0 mm SL; ZRC
38849, 1, 135.7 mm SL; Borneo: Kalimantan Barat: Ka-
puas drainage: Sungai Piyam. – ZRC 41947, 10, 40.4-
64.6 mm SL; Sumatra: Jambi.
Leptobarbus rubripinna: ZRC 49881, 2, 102.3-135.4 mm
SL; Thailand: aquarium trade. – CMK 17513, 6, 87.8-
92.4 mm SL; Thailand: Mae Khlong.
Additional material examined in the field.
Lobocheilos ixocheilos
Remarks. Described by Kottelat & Tan (2008).
Probably the species recorded from Jambi as
L. kajanensis by and Weber & de Beaufort (1916:
221).
Neobarynotus microlepis
Remarks. Not observed by us. Recorded by
Weber & de Beaufort (1916: 120). There is no
record of Neobarynotus collected after 1976 any-
where in its range, which originally included
Borneo, Sumatra, Malaysia, Cambodia and Viet-
nam (Kottelat & Widjanarti, 2005: 155).
Osteochilus kerinciensis, new species
(Figs. 30a-b)
Holotype. MZB 10711, 129.4 mm SL; Indonesia:
Sumatra: Kerinci, Danau Lingkat; H. H. Tan et
al., 10 June 1996.
Paratypes. ZRC 42288, 5; MZB 10712, 2; CMK
18914, 3; 115.1-151.0 mm SL; same locality as
holotype. – ZRC 42278, 2, 42.2-67.2 mm SL; Ker-
inci, Sungai Seli; H. H. Tan et al., 11 June 1996.
– ZRC 42284, 1, 92.0 mm SL; ZRC 43002, 3, 175-
215 mm SL; Kerinci, Danau Kerinci, near outflow
into Batang Merangi; H. H. Tan et al., 11 June
1996. – ZRC 43000, 1, 119.0 mm SL; Sungai
Dareh.
Diagnosis. Osteochilus kerinciensis shares with
O. waandersii and O. flavicauda the black midlat-
eral stripe ending at the posterior extremity of
the median caudal-fin rays, which distinguishes
them from all other species of the genus. Osteo-
chilus kerinciensis is distinguished from these two
species by the combination of: three tubercles at
tip of snout; red fins; transverse groove across
snout which appears as if pushed upwards; mouth
Tan & Kottelat: Fishes of Batang Hari
41
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
subinferior. See below for further characters
distinguishing these three species.
Description. General appearance in Figures 30a-b;
morphometric data of holotype and 5 paratypes
in Table 4. Dorsal fin with 3 simple and 11-12 {
branched rays; origin above lateral line scale 8-9.
Pectoral fin rounded to slightly angular, with 15
rays, reaching halfway to pelvic-fin base. Pelvic
fin triangular to slightly falcate, with 9 rays, reach-
ing about } of distance to anal-fin origin; axillary
scale present. Anal with 3 simple and 5 { branched
rays. Caudal fin with 10+9 principal rays, 9+8
branched. Caudal peduncle 1.41-1.65 times long-
er than deep. 31-32 + 2-3 scales along lateral line,
10-11 predorsal scales, { 5/1/6 { scales in trans-
verse line, { 3/1/3 { scales in transverse line on
caudal peduncle, 4 scales between lateral line and
pelvic origin.
Three tubercles at tip of snout, median one
much larger and conspicuous than lateral ones.
Main plicae on lateral portion of upper lip long
unbroken ridges (ectomorph type of Roberts, 1989:
46). Two pairs of maxillary barbels, length about
equal to eye diameter. About 39-45 gill rakers on
lower arm of first gill arch.
Coloration. Preserved: Body dark brown, paler
on belly. Dark and pale areas separated by a
midlateral stripe, from upper extremity of gill
opening to end of median caudal-fin rays, nar-
rower and fainter anteriorly, margins not discrete.
Pale area usually with conspicuous reticulated
pattern made of dark marks on scale pockets.
Head dark brown above; part below level of
lower eye margin pale yellowish brown; also
distinct on back, although fainter, in palest
specimens. Fins pale grayish brown.
Distribution. Osteochilus kerinciensis has been
Table 4. Morphometric data of holotype and 5 paratypes of Osteochilus kerinciensis.
holotype paratypes
MZB 10711 ZRC 42288 ZRC 43002
Standard length (mm) 129.4 144.0 135.5 212.0 169.8 169.5
In percent of standard length
Head length 22.6 22.3 22.5 21.8 22.0 22.7
Total length 133.5 131.7 134.4 137.8 132.5 134.1
Predorsal length 46.4 44.0 46.5 44.3 46.3 45.5
Prepelvic length 50.1 48.9 50.0 49.9 48.5 49.3
Preanal length 74.3 74.1 76.7 76.4 73.9 74.3
Head width 13.7 13.8 13.1 12.2 12.6 13.3
Head depth 16.5 16.9 16.3 16.0 16.3 16.8
Body depth at dorsal-fin origin 28.9 28.9 28.2 25.9 28.6 28.0
Body depth at anus 19.6 20.7 19.6 19.3 21.5 21.1
Depth of caudal peduncle 12.0 11.5 11.7 11.7 12.7 12.4
Length of caudal peduncle 17.5 18.0 19.4 17.7 19.6 17.6
Length of dorsal-fin base 28.4 27.3 24.8 27.1 26.7 27.3
Length of anal-fin base 10.2 10.4 10.3 9.5 10.9 9.7
Length of pelvic fin 20.4 19.1 18.3 18.6 19.1 19.1
Length of pectoral fin 19.1 18.4 18.7 17.4 17.9 18.3
Length of upper caudal lobe 34.5 33.8 37.2 39.8 36.6 34.7
Length of lower caudal lobe 33.3 31.9 33.7 38.7 33.5 33.3
Snout length 10.6 10.1 10.0 9.7 10.1 9.9
Eye diameter 4.5 4.2 4.6 3.5 4.1 4.0
Interorbital width 11.0 10.7 10.8 10.2 10.1 9.5
In percent of head length
Head depth 71 76 73 73 74 74
Head width 59 62 58 56 57 58
Snout length 46 45 45 44 46 44
Eye diameter 19 19 21 16 19 17
Interorbital width 47 48 48 47 46 42
42
Fig. 30. Osteochilus kerinciensis; Sumatra: Kerinci: Danau Lingkat; a, MZB 10711, holotype, 129.4 mm SL; b, about
150 mm SL, fresh, not preserved (June 1996).
collected in the upper part of the Batang Hari
drainage. The bulk of our material is from Lakes
Kerinci and Lingkat; a single specimen was ob-
tained at Sungai Dareh.
Etymology. Named for Lake Kerinci.
Remarks. Besides the characters mentioned in
the diagnosis, O. kerinciensis is distinguished from
O. waandersii (as discussed below) by: mouth
subinferior (vs. inferior), in ventral view, snout
slightly or not protruding beyond upper lip (vs.
conspicuously protruding); cheek appearing less
deep (not quantified, compare figures 30 and 34);
head shorter (21.8-22.8 % SL, vs. 23.0-25.8); snout
with blunt tip and transverse groove, appearing
as if pushed upwards (vs. deep, broadly rounded,
without transverse groove); and dorsal head
profile about as slanted as ventral one (vs.
clearly steeper).
Osteochilus kerinciensis differs from O. flavi-
cauda Kottelat & Tan, 2009 by: snout with a
transverse fold and tip pushed upwards (vs.
rounded, without transverse fold); fins red (vs.
yellow); three tubercles at tip of snout (vs. a sin-
gle tubercle); midlateral stripe faint, especially
anteriorly, with unsharp edges (vs. very con-
trasted throughout, with sharply marked edges);
and body with longitudinal rows of spots [pig-
ments on scale pockets] (vs. no longitudinal rows
of spots).
Osteochilus scapularis
Remarks. Our material agrees with Karnasutas
(1993: 31) and Roberts (1989: 47) descriptions of
O. enneaporos (Bleeker, 1852). But their descrip-
tions and illustrations do not agree with Bleekers
ones. Bleeker (1852c: 596) described the species
on the basis of a single specimen from Padang
(Sumatra); the holotype is figured natural size in
the Atlas (1863-64: 70, pl. 112 fig. 2). Our mate-
rial is distinguished from Bleekers in having a
much darker body, greater body depth (32.1-
37.2 % SL, vs. 28.4 on figure), very falcate dorsal
fin (when folded backwards, tip reaching beyond
base of last ray, vs. reaching base of 11th branched
ray [out of 12 {] ray), anal fin strongly falcate
(reaching caudal-fin base, vs. reaching about
halfway between base of last anal ray and caudal-
fin base), midlateral stripe faint, not contrasted
against background (vs. contrasted), snout blunt-
a
b
Tan & Kottelat: Fishes of Batang Hari
43
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
er, more rounded, and 3 tubercles at tip of snout
(vs. 9 main tubercles). Specimens over 150 mm
have the lower caudal-fin lobe shorter than the
upper and rounded, which probably results from
damages caused by the substrate (several rays
show signs of regrowth) (Fig. 31).
The holotype of O. enneaporos (Fig. 32) agrees
with the data in Bleekers text and figure and
most of the above characters are confirmed. Its
present length is 196 mm SL, the body depth is
28.9 % SL. The dorsal fin has 12 { branched rays;
the tip of the last simple ray is broken but reach-
Fig. 31. Osteochilus scapularis; Sumatra: Batang Hari drainage: Kiliran Jao; a, ZRC 49882, 184.4 mm SL; b, ZRC
49882, 184.4 mm SL, fresh (December 2003).
Fig. 32. Osteochilus enneaporos, BMNH 1866.5.2.172, holotype, 196 mm SL; Sumatra: Padang.
a
b
44
es about the base of the 10th branched ray. The
midlateral stripe is still faintly distinct on the
remaining lateral line scales of the overal discol-
ered specimen. Most scales are missing, but based
on the scale pockets it had about 32-34 + 2 lateral
line scales. The tubercles at tip of snout are miss-
ing but the sockets are clearly distinct. The 9 tu-
bercles reported by Bleeker in fact are the 3 large
tubercles observed in many species of Osteochilus
and two rows of 3 very small ones immediately
above and below.
At this stage, we have not yet been able to
identify any fresh material as Bleekers O. ennea-
poros. If the locality data (Padang) is accurate,
then this is not surprising as the numerous small
coastal drainages of the Indian Ocean slope of
Sumatra seem to be inhabited by their own en-
demics and they have not yet been surveyed (or
the results remain unpublished).
Our material agrees with the original descrip-
tion and figure of O. scapularis Fowler, 1939, which
had been treated as a synonym of O. enneaporos
by Karnasuta (1993) and Roberts (1989). Fowlers
(1939) figure and description show a deep-bodied
fish, with a high, rounded snout, a midlateral
stripe distinct only posteriorly and long and
falcate dorsal and anal fins. We treat O. scapularis
as the valid name of the species called O. ennea-
poros by Karnasuta & Roberts. It is distributed in
Malay Peninsula (from Trang southwards), Bor-
neo (Kapuas, Sarawak) and Sumatra (Batang
Hari).
Osteochilus vittatoides Popta, 1904 (see also
Popta, 1906: 94) (Fig. 33), originally described
from the Mahakam and Kayan drainages in east-
ern Borneo, has been treated as a synonym of
O. enneaporos by Karnasuta (1993: 31) and Roberts
(1989: 47) (our O. scapularis). Poptas original
description largely disagrees on a number of
points with our material of O. scapularis. Recent
material from the Kayan agrees well with Poptas
description and we can confirm that O. vittatoides
differs from O. scapularis as follows: dorsal fin
concave (vs. very falcate), first branched ray
reaching at most base of last one (vs. reaching
beyond), more slender body (depth 27.2-29.8 %
SL, vs. 32.1-37.2), several very small tubercles on
tip of snout (vs. 3 large tubercles), shorter snout
(10.3-11.6 % HL, vs. 11.1-13.6; 44-49 % HL, vs.
47-53), more slender caudal peduncle (depth 11.5-
12.7 % SL, vs. 13.5-14.8; depth 1.35-1.65 times in
its length, vs. 1.11-1.36), tip of first branched anal-
fin ray not reaching base of caudal fin (vs. reach-
ing beyond), all fins pale yellowish brown (vs.
orange to red), and black midlateral stripe faint
and inconspicuous in large adults, and with ir-
regular edges (vs. distinct and with more discrete
edges). While all examined O. vittatoides have 11 {
branched dorsal-fin rays, O. scapularis has 11-13 {.
Osteochilus vittatoides is a valid species restricted
to eastern Borneo drainages.
Osteochilus vittatoides is distinguished from
O. enneaporos as described by Bleeker (1852c) by:
having the lateral stripe extending to tip of me-
dian caudal rays (vs. ending at caudal-fin base),
11½ branched dorsal-fin rays (vs. 12 {), yellowish
brown fins (vs. reddish-orange).
Material. Osteochilus enneaporos: BMNH 1866.5.2.172,
holotype, 196 mm SL; Sumatra: Padang.
Osteochilus scapularis: ZRC 49882, 3; CMK 20839, 2;
91.9-184.4 mm SL; Sumatra: Sumatera Barat: Kiliran Jao,
H. H. Tan et al., Dec 2003.
Osteochilus vittatoides: ZRC 49883, 11; CMK 18916,
5; 64.5-118.7 mm SL; Borneo: Kalimantan Timur: Kayan
drainage, Sungai Mutai; H. H. Tan et al., Nov 1999.
Osteochilus vittatus
Remarks. Rohita vittata Valenciennes (in Cuvier
& Valenciennes, 1842: 267) and R. hasseltii Valen-
ciennes (in Cuvier & Valenciennes, 1842: 274) are
simultaneous subjective synonyms. The species
has long been known as O. hasseltii (at least since
Weber de Beaufort, 1916) and therefore Kottelat
(1989: 9) gave precedence to R. hasseltii. Unfortu-
nately, there is an earlier first reviser act: Guibé
& Spillmann (1958: 463) had already realised the
synonymy and given precedence to R. vittata.
Therefore the valid species name is O. vittatus.
Weber & de Beaufort (1916: 139) recorded
O. kappenii (Bleeker, 1857) from Jambi. This species
is otherwise known only from the Kapuas drain-
age, where it replaces O. vittatus (Roberts, 1989:
49; Karnasuta, 1993: 66; Kottelat & Widjanarti,
2005: 151). The record from the Baram (Sarawak)
by Fowler (1905) refers to O. kahajanensis (see
Karnasuta, 1993: 66). We have not seen O. kap-
penii in the Batang Hari drainage and we do not
include it in our list and we hypothesise that it is
a misidentification of O. vittatus.
Osteochilus waandersii
Remarks. The Batang Hari collections include
two species which key out as O. waandersii using
Karnasutas (1993) key. They were obtained in
the foothills and highlands. Fresh specimens of
Tan & Kottelat: Fishes of Batang Hari
45
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
both have a golden to yellowish body with a
conspicuous black lateral line reaching to the end
of the median caudal-fin rays and orange to red-
dish fins. One of them is identified as O. waander-
sii and the other, distinguished by its shorter head
and with different habitat preferences, is de-
scribed above as O. kerinciensis.
Bleeker (1852d: 733) described O. waandersii
on the basis of a single specimen 191 mm TL from
Bangka, which he figured natural size in the
Atlas (Bleeker, 1863-64: 63, pl. 110 fig. 2). Al-
though there are slight differences, the specimens
we identify as O. waandersii largely agree with
Bleekers holotype, figure and description. The
holotype (BMNH 1866.5.2.169; Fig. 34a) has a
pointed and less deep snout (vs. blunt in our fresh
material; but the snout is less pointed in the ac-
tual holotype than on Bleekers figure); { 6 scales
between the dorsal-fin origin and the lateral line
(vs. { 5); the cheek appears less deep (not quan-
tifiable, but compare Figures 34b-c). Although the
head of our material appears longer than the head
of the holotype figured by Bleeker (23.0-25.8 %
SL, vs. 21.6), we measure it as 23.9 % SL in the
actual holotype.
Our attempts to collect O. waandersii on Bang-
ka Island (the type locality) failed. Out of the two
Batang Hari species, O. kerinciensis is presently
known only from the upper part of the drainage
and Lake Kerinci; O. waandersii was found in the
foothills and is also known throughout the Sunda
Shelf around Bangka (the Malay Peninsula, Bor-
neo and Java; see, i.a., Kottelat et al., 1993: pl. 13),
in similar habitats and we presently have no
reason to doubt that the Bangka holotype is con-
specific.
Bleeker (1863-64: 68, pl. 113 fig. 2) recognized
a second species of Osteochilus with a conspicuous
black midlateral stripe extending to the end of
the median caudal-fin rays under the name Ro-
hita vittata, a species originally described by Va-
lenciennes (in Cuvier & Valenciennes, 1842).
Karnasuta (1993: 81) treated R. vittata as a junior
synonym of O. hasseltii (see also Kottelat, 1989: 9,
see also above). He further commented that O. vit-
tatus of Bleeker is mis-identified and in fact is
O. microcephalus, a point on which we disagree.
Bleekers plate unambiguously shows the black
midlateral stripe reaching the posterior extrem-
ity of the median caudal rays (as in our material)
while it does not extend on the caudal fin in
O. microcephalus. Bleekers figure of O. vittatus
also shows the deep snout and the anterior ex-
tremity of the midlateral stripe at upper extrem-
ity of the gill opening and at level of upper
margin of eye, characters diagnostic of O. waander-
sii as recognized here. We note, however, some
variability within the Sundaic material that we
identify as O.
waandersii. Some specimens (includ-
ing the holotype and the Batang Hari material)
are relatively deeper-bodied and with a less con-
trasted stripe, while others have a more slender
and more yellowish body with a more contrasted
stripe, more closely resembling the figure of
R. vittata in Bleeker (1863-64: pl. 113 fig. 2) (see
figure in Kottelat et al., 1993: pl. 13). This suggests
that two species might be involved, but we are
unwilling to be definitive on this issue before
seeing topotypes of O. waan dersii.
In the attempt to clear the identity of O. waan-
dersii, we examined various material from the
Malay Peninsula earlier identified as O. waander-
sii, O. microcephalus yellow fins or O. enneaporos
yellow fins (e.g., Zakaria-Ismail, 1987; Ng & Tan,
1999). This materials is described elsewhere as
O. flavicauda Kottelat & Tan, 2009. The specimen
from Pattani (Thailand) illustrated as O. waander-
sii by Karnasuta (1993: 30, fig. 13) apparently
belongs to O. flavicauda, as does the fish figured
by Smith (1945: 216, pl. 6) as O. vittatus. The ma-
terial from the Mekong drainage identified as
O. waandersii by Kottelat (1998: 45; 2001: 66)
should now be compared with the Sundaic mate-
rial. Should it turn out to represents a distinct
species, the name O. soplaoensis (Fowler, 1934) is
available for it.
Material. BMNH 1866.5.2.169, holotype, 152 mm SL;
Indonesia: Bangka: Toboali Prov. – ZRC 49884, 1,
204 mm SL; Sumatra: Sumatra Barat; Kiliran Jao. – ZRC
45535, 8, 32.8-90.0 mm SL; Kalimantan Timur: Kayan.
– ZRC 38886, 1, 64.1 mm SL; CMK 11865, 4, 56.0-72.3 mm
SL; Kalimantan Tengah: Barito. – ZRC 42782, 1, 127.3 mm
SL; Malaysia: Johor: Sungai Kahang. – ZRC 42642, 6,
51.8-112.6 mm SL; Malaysia: Pahang: Sungai Kinchin.
– ZRC 1889, 1, 183 mm SL; Malaysia: Perak: Bukit Meh-
ra Reservoir. – ZRC 41861, 1, 131.3 mm SL; CMK 8130,
1, 49.9 mm SL; Malaysia: Terengganu: Sungai Brang.
Pectenocypris micromysticetus, new species
(Figs. 35a-c)
Holotype. MZB 10993, 43.0 mm SL; Sumatra:
Jambi: Danau Semangkat, a lake connected to
Batang Hari by Sungai Bangko, opposite Kam-
pung Senaning; M. Kottelat, 6 June 1994.
46
Fig. 33. Osteochilus vittatoides, ZRC 49883, 118.7 mm SL; Borneo: Kayan drainage (right side, reversed).
Fig. 34. Osteochilus waandersii; a, BMNH 1866.5.2.169, holotype, 152 mm SL (right side, reversed); Bangka; b, ZRC
42642, 103.7 mm SL; Malaysia: Pahang: Ulu Kinchin; c, ZRC 49884, 204 mm SL; Sumatra: Batang Hari drainage,
Kiliran Jao, fresh (December 2003).
b
a
c
Tan & Kottelat: Fishes of Batang Hari
47
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 35. Pectenocypris micromysticetus; a, MZB 10993, holotype, 43.0 mm SL; Sumatra: Batang Hari drainage,
Danau Semangkat; b, ZRC 51759, fresh, about 36 mm SL, Sumatra: Batang Hari drainage: Danau Arang Arang
(July 1997); c, ZRC 38675, 41.7 mm SL, opercle removed to show gill rakers (right side, reversed).
Paratypes. ZRC 38675, 48; MZB 6001, 10; CMK
11248, 40; 31.8-44.9 mm SL; same data as holotype.
– ZRC 51758, 46, 31.7-43.2 mm SL; Sumatra:
Jambi, Pijoan, Danau Souak Padang; H. H. Tan
et al., 8 Jun 1996.
Additional material (non types). ZRC 38670, 27; CMK
11242, 27; 14.4-22.6 mm SL; Sumatra: Jambi: Danau Kiri,
a lake connected to Batang Hari by Sungai Bangko,
opposite Kampung Senaning; M. Kottelat, 6 June 1994.
– ZRC 38532, 11; CMK 11092, 10; 11.5-25.5 mm SL;
Sumatra: Jambi: Danau Arang Arang; M. Kottelat & H.
H. Tan, 29 May 1994.
Diagnosis. Pectenocypris micromysticetus is dis-
tinguished from P. korthausae Kottelat, 1982 in
having more gill rakers on the anterior gill arch
(45-70 + 120-170, vs. 20-35 + 75-100; see also Re-
marks) and an elongate blotch at middle of caudal-
fin base and end of caudal peduncle (vs. rounded
to triangular, occupying {-} of depth of caudal
peduncle). It shares the high number of gill rakers
with P. balaena Roberts, 1989, from which it is
distinguished by the absence of or only very faint
midlateral stripe on the body (vs. presence) and
the presence of a conspicuous, elongated black
b
a
c
48
blotch at caudal-fin base (vs. no blotch or only a
small blackish dot terminating the mid-lateral
stripe).
Description. See Figs. 35a-b for overall appear-
ance, and Table 5 for morphometric data of holo-
type and 9 paratypes. Body slender, compressed.
Body depth about equal to head depth. Dorsal
outline of body slightly arched; ventral outline
straight to slightly convave between head and
anus, then slanted and straight to caudal-fin base.
Mouth terminal, lower jaw slightly projecting.
Gill rakers long, very thin, very densely set and
difficult to count with accuracy. Counted in 4
specimens: ZRC 38675, 44.9 mm SL, about 70+170;
ZRC 38675, 41.7 mm SL (Fig. 35c), about 67+160;
ZRC 38675, 32.5 mm SL, about 55+150. In ZRC
38532, 25.4 mm SL, at least 45+120. Number of
gill-rakers possibly increasing with SL.
Dorsal fin with 2 simple and 6 or 7 branched
rays; last ray short (not easily distinguishable),
slender, not branched, not articulating on same
pterygiophore as ray 6; origin above pelvic-fin
origin. Pectoral fin short, rounded, reaching about
} of distance to pelvic-fin base, with 12-14 rays;
no axillary lobe. Pelvic fin triangular, slightly
pointed, reaching to or almost to anus, with 8
rays; axillary scale present, not significantly
longer than other scales. Anal with 3 simple and
5½ branched rays; posterior edge concave. Caudal
fin deeply forked, with pointed lobes, with 10+9
principal rays, 9+8 of them branched. Caudal
peduncle 1.8-2.1 times longer than deep.
Scales fragile, many missing in preserved
specimens. Lateral line incomplete, perforating 7
or 8 scales; lateral line row scales 28-29 + 2-3. 11-12
predorsal scales, { 8 { scales in transverse line
(counted 2 scale rows in front of pelvic-fin base),
{ 5 { scales in transverse line on caudal peduncle,
1 scale between lateral line row and pelvic-fin
origin.
Coloration. After fixation in formalin and storage
in ethanol. Body yellowish brown, darker on
dorsal half. Somewhat darker in middle of body,
resulting in a very vague stripe, sometimes dis-
tinct on caudal peduncle, margined above by a
paler stripe (after storage in ethanol, midlateral
area of anterior part of body appears darker but
this is mostly an artefact; because of dehydration
due to ethanol, a pleat is formed in this area and
the shade appears as a stripe). A thin black axial
streak from caudal-fin base forwards until verti-
cal of dorsal fin. A thin black mid-dorsal stripe
from head to caudal-fin base. A thin black stripe
from anus to caudal-fin base, running along anal-
fin base and mid-ventral outline of caudal pedun-
cle. Head yellowish brown, darker dorsally, with
an unpigmented area forming a semi-circular
patch around lower half of eye.
A conspicuous black blotch at posterior ex-
tremity of caudal peduncle and caudal-fin base,
elongate, about twice longer than deep, occupy-
ing about ~ of depth of caudal peduncle. Blotch
surrounded by a paler, reddish area, more con-
spicuous above and below. Fins hyaline, with
scattered black pigments in proximal areas of
rays, and along edges of rays.
In life. Body greyish brown. An iridescent
stripe above axial streak, between gill opening
and caudal blotch.
Distribution. Pectenocypris micromysticetus is
presently known only from the Batang Hari drain-
age. It is common in oxbow lakes and still pools
in open or recently flooded areas.
Etymology. From the Greek μικρος (mikros),
small, μυ
σταξ (mustax), moustache, and κη
τος
(kêtos), sea monster (by extension, the Latin cetus,
Table 5. Morphometric data of holotype (MZB 10993)
and 9 paratypes (ZRC 38675) of Pectenocypris micromys-
ticetus. Holotype values included in range. H, holo-
type.
H range mean
Standard length (mm) 43.0 40.7-44.9
Total length (mm) 55.3 53.1-57.3
In percent of standard length
Head length 27.8 26.4-28.9 27.6
Predorsal length 52.7 52.2-55.7 53.9
Prepelvic length 51.8 51.7-56.0 53.4
Pre-anal length 72.7 71.9-76.3 73.8
Head depth 16.0 15.3-16.6 16.0
Body depth at dorsal-fin origin 17.9 17.8-19.2 18.3
Depth of caudal peduncle 10.6 9.8-10.7 10.2
Length of caudal peduncle 20.7 19.0-21.2 20.2
Snout length 8.0 6.8-8.7 7.8
Eye diameter 8.0 7.2-8.6 7.8
Interorbital width 9.4 8.7-9.4 9.0
Dorso-hypural distance 49.0 45.1-49.0 47.0
In percent of head length
Snout length 29 24-31 28.1
Eye diameter 29 26-31 28.2
Interorbital width 34 30-34 32.7
Tan & Kottelat: Fishes of Batang Hari
49
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
large sea-animal, gave rise to the biological word
ceatacea, the whales). Allusion to the numerous
gillrakers appearing like baleen plates of mysticete
whales. A noun in apposition.
Remarks. We have several samples of small-size
Pectenocypris, which we tentatively refer to P. mi-
cromysticetus. The gill-rakers are very difficult to
count, but specimens around 15-20 mm SL have
about 70-120 gill rakers on the upper limb of the
first gill arch (ZRC 38670, CMK 11242). The data
reported above suggest that there is some increase
of the number of gill rakers with size, but it is not
clear if this increase could be as large as from
about 70-120 at 15-20 mm SL to about 150-170 at
32-45 mm SL.
The lowest gill-raker counts are within the
range known in larger, adult P. korthausae (20-35 +
75-100 at 20-30 mm SL; Kottelat, 1982; Roberts,
1989; pers. obs.) but these specimens are not
P. korthausae. Compared side by side with topo-
types of P. korthausae of about similar size they
have a different head and body shape, with the
head depth at nape greater than depth at dorsal-
fin origin (vs. about identical or somewhat
smaller in P. korthausae), body outline between
head and pelvic-fin base slightly concave or
straight (vs. slightly convex), a more rounded
dorsal outline of head, and a stouter caudal pe-
duncle (deeper; not quantified). Subjectively,
these specimens give the impression of juveniles
while at a similar size (about 18-22 mm SL) P. kort-
hausae gives an adult impression. To date,
P. korthausae is not known to exceed 35 mm SL
(45 in P. micromysticetus); it is known only from
Kalimantan Tengah.
Comparison material. Pectenocypris balaena: ZRC 51757,
49, 26.5-37.5 mm SL; Borneo: Kalimantan Barat: Kapuas
drainage: Lake Sentarum area: Danau Lebuyan.
Pectenocypris korthausae: ZRC 22856, 5, 21.1-29.3 mm
SL; Borneo: Kalimantan Tengah: Mentaya drainage. –
ZRC 51755, 51; CMK 20819, 52; 22.2-27.2 mm SL; Borneo:
Kalimantan Tengah; Kahayan drainage.
Puntius sp. Bertam
Remarks. A member of the P. binotatus group,
known from Sumatra and Malay Peninsula (Kot-
telat & Lim, in prep.).
Puntius sp. Kerinci
Remarks. A member of the P. binotatus group,
endemic to the Indragiri and Batang Hari drain-
ages, found primarily in volcanic lakes Danau di
Atas, Gunung Tujuh and Kerinci (Kottelat & Lim,
in prep.). This species and Homaloptera gymno-
gaster are the highest altitude fish in Sumatra,
found in Danau Gunung Tujuh at 1950 m a.s.l.
Raiamas guttatus
Remarks. New record for Sumatra (see Fig. 36).
Restricted to headwaters and foothill rivers.
Material. ZRC 51776, 1, 212.0 mm SL; Sumatra: Su-
matra Barat, Kiliran Jao. – ZRC 51784, 2, 86.4-126.0 mm
SL; Sumatra: Sumatra Barat, Sungai Dareh.
Rasbora brittani
Remarks. New record for Sumatra and first
record since original description. The species was
based on aquarium specimens, reportedly from
Johor River at Segamat, Malaysia (Axelrod, 1976),
where, despite several attempts, we have not been
able to find it. We cannot exclude that these local-
ity information are erroneous.
The generic placement of this species requires
confirmation. It is missing the symphysal knob
in the lower jaw, which is otherwise known in all
other rasborines.
Rasbora sumatrana
Remarks. This species is found at the foot of
Gunung Tujuh and is the species of Rasbora found
in the highest altitude in Sumatra (Fig. 37). As
re-diagnosed in Kottelat et al. (1993: 48 [65],
pl. 20), R. sumatrana is restricted to the fast flow-
ing streams of the interior of Sumatra. It is re-
corded only from the Batang Hari and Indragiri
drainages. Even so, the identity of the various
populations of R. sumatrana is not yet clear. We
observe variability between populations, but the
number of populations available for study is too
small to reach conclusions and we tentatively
retain all in the same species. The type locality is
Solok, in the Indragiri drainage.
The name R. sumatrana has been used for a
variety of species in Southeast Asia. The Indochi-
nese R. sumatrana are now recognised as R. pa-
viana Tirant, 1885 (see Kottelat, 2001), those from
the Malay Peninsula as R. vulgaris Duncker, 1904
and R. notura Kottelat, 2005. Examination of fresh
50
Fig. 36. Raiamas guttatus, ZRC 51776, 212.0 mm SL, fresh (December 2003); Sumatra: Batang Hari drainage, Kiliran
Jao.
Fig. 37. Rasbora sumatrana; a, ZRC 42289, 57.0 mm SL; Sumatra: Batang Hari drainage, Danau Lingkat; b, CMK
4642, 66.6 mm SL; Sumatra: Indragiri drainage: Solok.
material from Sarawak and Kalimantan Barat
shows that they differ from R. sumatrana s.s. as
follow: the pelvic-fin origin is separated from the
lateral line scale row only by the axillary scale
(vs. by one scale row plus the axillary scale); the
tip of the anal fin is pointed (vs. blunt); the inter-
orbital area is almost flat (vs. convex in R. suma-
trana); in lateral view, the dorsal profile of the
head is straight to slightly concave, with the tip
of the snout often slightly turned upwards (vs.
convex, with the snout pointed forwards); the
midlateral stripe is often incomplete (when com-
plete then usually not very contrasted, or faint
anteriorly), running below the axial streak and
separated from it by a pale area at the level of the
dorsal-fin origin, and ending in a faint widened
b
a
Tan & Kottelat: Fishes of Batang Hari
51
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
patch at the end of the caudal peduncle (vs. begin-
ning above the pelvic-fin origin, running along
or over the axial streak for all its length, ending
in a small vertically elongated black dot at caudal-
fin base); the scales below the midlateral stripe
have a dark pocket and a black edge forming a
reticulated pattern, but they are not covered by
pigments (vs. at least half of the exposed part of
the scales immediately under the anterior extrem-
ity of the midlateral stripe dark brown, sometimes
even appearing as a diffuse brownish blotch); the
caudal fin has a black margin and in many
populations the tip of the lobes is black (vs. no
black marking on caudal fin); and 17-18 + 15-17
= 33-35 vertebrae (vs. 17 + 14-16 = 31-33 [Kottelat
& Vidthayanon, 1993]).
In most aspects, the Sarawak R. sumatrana
agrees with the description of R. hosii Boulenger,
1895, but it disagrees in having the pectoral fin
reaching the pelvic fin (vs. not) and 24-25 + 2
lateral line scales (vs. 28-29). We have examined
photographs and radiographs of the syntypes of
R. hosii (BMNH 1894.8.3.65-66). The 80.2 mm SL
(BMNH 1894.8.3.65) syntype (here designated as
lectotype; Fig. 38a) shares all the diagnostic char-
acters of our Sarawak R. sumatrana, except for
the colour marks, which are all faded or missing
on the broken caudal fin. Some scales still have
the distinct black posterior edge. The pectoral fin
is broken but was probably not reaching the
pelvic-fin base. There are 26+2 lateral line scales.
The 86.3 mm SL paralectotype is less well pre-
served and more scales are missing. We conclude
that R. hosii is the valid name of the Sarawak
R. sumatrana (Fig. 38b). The type locality of
R. hosii is the Baram River (Sarawak) and we have
examined material from adjacent drainages in
Brunei. The Sabah material listed by Kottelat &
Vidthayanon (1993) as R. cf. sumatrana is an un-
named species (Kottelat & Tan, in prep.).
Material. Rasbora hosii: ZRC 39379, 2, 49.9-61.9 mm SL;
Borneo: Sarawak: Kuap (1°25'08.9"N 110°20'52.9"E). –
CMK 10920, 2; Borneo: Sarawak: Sungai Semabang. –
ZRC 42725, 20, 19.1-76.9 mm SL; Borneo: Brunei Darus-
Fig. 38. Rasbora hosii; a, BMNH 1894.8.3.65, lectotype, 80.2 mm SL; Borneo: Sarawak: Baram River (copyright:
Natural History Museum, London); b, CMK 10920, 48.2 mm SL; Borneo: Sarawak: Sungai Semabang.
b
a
52
salam: Belait District: Sungai Ingei. – ZRC 38826, 6,
36.9-62.2 mm SL; CMK 11685, 25, 33.6-75.5 mm SL;
Borneo: Kalimantan Barat: Sungai Lanjak. – ZRC 39385,
15, 43.0-74.7 mm SL; Borneo: Sarawak: Matang: Sungai
Cina Matang.
Rasbora sumatrana: ZRC 42289, 20, 16.6-58.4 mm SL;
Sumatra: Kerinci, Danau Lingkat. – ZRC 42279, 4, 37.9-
45.6 mm SL; Sumatra: Kerinci: Sungai Seli. – ZRC 42273,
2, 22.7-42.1 mm SL; Sumatra: Kerinci: Sungai Jalnei
Dalam, at base of Gunung Tujuh. – CMK 4642, 10;
Sumatra: Sumatera Barat: Solok.
Schismatorhynchos heterorhynchos
Remarks. Schismatorhynchos heterorhynchos is
restricted to headwater and foothill rivers in
Sumatra (Indragiri, Batang Hari, Musi) and Ka-
limantan Barat (Upper Kapuas) (Siebert & Tja-
krawidjaja, 1998; pers. obs.). The juveniles have
a violet sheen on the body with reddish fins (Fig.
39a) and adults are dark purple-grey with grey
fins (Fig. 39b).
Tor tambroides
Remarks. Tor tambroides is cultivated in Kerinci
(apparently grown up in cages from wild caught
juveniles) and is a popular food fish, but local
populations have declined due to overfishing.
Trigonostigma hengeli
Remarks. Trigonostigma heteromorpha (Duncker,
1904) is found from the Indragiri drainage north-
wards and T. hengeli is known from the Batang
Hari drainage and possibly occurs further south.
Trigonostigma hengeli was recently obtained from
the area of Pontianak, Kalimantan Barat, Borneo
(Kottelat & Witte, 1999). We have examined ma-
terial of all named species and all known popula-
tions of Trigonostigma and for the time being retain
T. hengeli as the name of the species occuring in
the Batang Hari drainage in Sumatra and the
coastal peat swamps of Kalimantan Barat.
Family Balitoridae
Homaloptera gymnogaster
Remarks. Highland species relatively common
in riffles and quiet waters of Sumatra. Highest
altitude balitorid in Sumatra, found at 1950 masl
in Danau Gunung Tujuh.
Neohomaloptera sp.
Remarks. Differs from N. johorensis chiefly in
body coloration and longer nasal barbel. The
Batang Hari collections include a single specimen
14.9 mm SL from Sungai Ayer Merah, near Pi-
joan.
Family Nemacheilidae
Nemacheilus cf. kapuasensis
Remarks. Obtained only from upstream area.
We have only one sample of five specimens in
poor state that we tentatively refer to N. kapuas-
ensis. They do not belong to any of the nominal
species of the genus known from Sumatra. The
identity of N. jaklesii (Bleeker, 1852c) is still not
clear, but the present specimens do not belong to
N. jaklesii. Bleeker (1852c) described N. jaklesii
with 11 or 12 bars and in 1863-64 he figured it
with 12 very narrow and widely spaced bars,
while the present specimens have 11-13 bars
wider than the interspaces or a row of saddles
along the back more or less connected to a row
of blotches along the lateral line.
Material. ZRC 41561, 5, 48.7-54.3 mm SL; Sumatra:
Sumatera Barat: Sungai Dareh, Pulau Punjung market
(bridge – 0°57'53.9"S 101°30'18.9"E).
Nemacheilus longipinnis
Remarks. The Batang Hari collections include a
single specimen with totally white body except
for a faint midlateral stripe and a black spot at
middle of caudal-fin base. This is the colour pat-
tern of large individuals of N. lactogeneus Roberts,
1989 as figured by Kottelat et al. (1993: pl. 25).
The original description and illustration do not
mention or show the faint stripe, but as the mate-
rial figured by Kottelat et al. (1993) comes from
the same area (Kapuas River upstream of Sintang)
we consider this as intraspecific variation. We
also have seen material conspecific with the Bor-
nean one from Sumatra (unfortunately with un-
precise locality data: Seberida, either in Indragiri
or Kampar Kiri drainages).
Nemacheilus longipinnis Ahl, 1922 was de-
scribed from central Sumatra (without precise
locality data) on the basis of a single, poorly
preserved specimen without any colour pattern
(Kottelat et al., 1993: 55 [76], fig. 139). The species
has not been reported since (see Kottelat, 1984).
Tan & Kottelat: Fishes of Batang Hari
53
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
The holotype is noteworthy for its very large eye
(15 times in SL, 3.2 times in HL). In our specimen,
the eye diameter is 15.7 times in SL and 3.3 times
in HL (SL 34.5 mm, HL 7.3 mm, eye diameter
2.2 mm) and with the limited available material
we identify it as N. longipinnis. In the three Se-
berida specimens the eye diameter is 15.1-19.8
times in SL and 3.6-3.9 times in HL and we con-
clude that they are conspecific with the Jambi
specimen. Roberts (1989: 107) records that the eye
diameter is 18.9-19.8 times in SL for the two types
of N. lactogeneus. Our only three adult and sub-
adult specimens from the Kapuas drainage have
the eye diameter 14.6-18.4 times in SL and 3.4-4.2
times in HL. The comparison of our small samples
suggests that the Kapuas material too (N. lactoge-
neus) might be conspecific with N. longipinnis. The
only difference we observed is that the eyes are
slightly more dorsally located eyes and the inter-
orbital area is flat (vs. slightly convex) in the
Kapuas material.
Material. ZRC 42303, 1, 34.5 mm SL; Sumatra: Jambi:
Sungai Alai. – CMK 8354, 3, 51.5-61.2 mm SL; Sumatra:
Riau: vicinity of Seberida. – CMK 6890, 37, 13.2-53.4 mm
SL; Borneo: Kalimantan Barat: Kapuas River mainstream
at Teluk Ujung Bayur. – CMK 10462, 5, 16.7-51.8 mm
SL; Borneo: Kalimantan Barat: Sungai Kapuas at Nanga
Al-Fazal.
Nemacheilus papillos, new species
(Figs. 40-41)
Holotype. MZB 10994, 55.3 mm SL; Sumatra:
Sumatera Selatan: Sungai Sentang near Desa
Sukajaya, about 5 km from road (turn-off at about
12 km on road from Bayung Lencir to Jambi; H. H.
Tan, 27 July 1997.
Paratypes. ZRC 43096, 6, 47.3-55.7 mm SL; ZRC
51760, 1, 56.5 mm SL; same data as holotype. –
ZRC 42398, 15, 17.6-40.2 mm SL; CMK 13019, 4,
37.5-39.1 mm SL; Sumatra: Jambi: Sungai Pijoan
main confluence; before entry in Danau Souak
Table 6. Morphometric data of holotype (MZB 10994) and 4 paratypes (ZRC 43096) of Nemacheilus papillos.
holotype paratypes
Standard length (mm) 55.3 55.7 52.4 52.8 50.4
In percent of standard length
Total length 133.1 136.4 135.9 134.1 134.9
Head length 20.1 20.1 19.8 20.3 25.0
Predorsal length 47.0 47.9 46.6 47.3 48.4
Prepelvic length 49.9 50.1 49.4 50.6 49.0
Preanal length 76.5 78.3 76.3 78.2 77.8
Head width 13.7 13.1 12.8 13.1 13.9
Body depth at dorsal-fin origin 14.5 15.3 15.3 20.3 14.3
Body width at dorsal-fin origin 9.8 10.1 11.3 9.5 9.1
Depth of caudal peduncle 11.4 12.0 12.2 11.2 12.1
Length of caudal peduncle 12.8 16.2 13.9 13.1 14.9
Length of base of dorsal fin 20.6 21.2 22.1 19.7 21.8
Length of base of anal fin 8.5 10.4 11.1 11.6 11.3
Length of pelvic fin 15.9 17.8 18.1 18.4 17.7
Length of pectoral fin 25.5 22.6 23.5 22.7 26.0
Length of upper caudal lobe 35.4 33.9 38.2 34.7
Length of median caudal rays 19.9 22.4 18.1 18.4 17.7
Length of lower caudal lobe 32.0 34.2 33.7 34.7
Snout length 8.9 9.3 9.2 9.1 9.3
Eye diameter 5.2 5.2 4.6 4.7 5.4
Interorbital width 6.7 7.0 6.5 6.4 6.9
In percent of head length
Head width 69 65 64 65 56
Snout length 44 46 46 45 37
Eye diameter 26 26 23 23 21
Interorbital width 33 35 33 32 28
54
Padang; H. H. Tan et al., 8 Jun 1996. – ZRC 43034,
14, 20.6-28.1 mm SL; Sumatra: Jambi: Leibong
Sepbaju, stream in swampforest; H. H. Tan, 21
Nov 1996.
Diagnosis. Nemacheilus papillos is distinguished
from the other species of the genus on the Sun-
daic area by having strongly papillated lips
(Fig. 41), the anterior nostril at the tip of a short
tube and the colour pattern consisting in 10-11
bars, much wider at level of lateral line, bars
behind dorsal-fin origin fused at level of lateral
line. The following characters are useful for iden-
tification, although none is unique to the species:
no acuminated scales on caudal peduncle; black
bar across eye; black blotch at base of anterior
dorsal-fin rays; caudal forked, upper lobe longer
than lower one; no sexual dimorphism; caudal-fin
with 9+8 branched rays.
Description. See Figure 40 for general appear-
ance and Table 6 for morphometric data of holo-
type and 4 paratypes. A moderately elongate
nemacheiline with body depth about equal from
nape to caudal-fin origin, only sligthly tapering
on caudal peduncle. Interorbital area flat, in lat-
Fig. 39. Schismatorhynchos heterorhynchos; Sumatra: Batang Hari drainage, Kiliran Jao; a, ZRC 51777, about 100 mm
SL, fresh (Dec 2003); b, ZRC 51777, about 240 mm SL, fresh (Dec 2003).
b
a
Tan & Kottelat: Fishes of Batang Hari
55
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
eral view eye flushed with dorsal profile of head.
Head slightly compressed; body slightly com-
pressed anteriorly to compressed posteriorly.
Pectoral fin reaching or almost reaching pelvic-fin
base. Axillary pelvic lobe present. Pelvic fin almost
reaching anus, which is about one eye diameter
in front of anal-fin origin; origin under base of
branched ray 2. Caudal fin forked, upper lobe
clearly longer than lower one. Low ventral and
dorsal adipose crests at posterior extremity of
caudal peduncle which is 1.18-1.31 times longer
than deep. Distal margin of dorsal fin straight.
Largest recorded size 56 mm SL.
D 4/9 {. A 3/5 {. C /9+8/. P 15. V 7.
Body covered by small embedded scales,
except belly in front of pectoral-fin base. Lateral
line complete, with about 94-111 pores (difficult
to count with accuracy). Cephalic lateral line
system with 6 supraorbital, 4+12 infraorbital, 9-10
preoperculo-mandibular and 3 supratemporal
pores.
Fig. 40. Nemacheilus papillos, MZB 10994, holotype, 55.3 mm SL; Sumatra Selatan: Sungai Sentang (right side,
reversed).
Fig. 41. Nemacheilus papillos, ZRC 43096, 54.1 mm SL;
mouth. Scale bar 1 mm (drawing by Kelvin Lim).
56
Anterior nostril at tip of a short tube. Mouth
gape about 2 times wider than long (Fig. 41). Both
lips strongly papillated. A narrow median inter-
ruption in lower lip. Processus dentiformis
present, pointed. No median notch in lower jaw.
Inner rostral barbel reaching vertical through
middle of eye; both outer rostral and maxillary
barbels reaching about midway between eye and
posterior margin of opercle. Intestine with a loop
immediately behind stomach.
Sexual dimorphism. None observed.
Coloration. Colour pattern is shown on Figure
40. Body yellowish to light brown. Ten to 11 dark
brown bars on body, wider than interspaces,
wider at level of lateral line, bars behind vertical
of dorsal-fin origin fused on lateral line, anterior
bars sometimes vertically split below lateral line;
bars joining across dorsal midline, not joining
across ventral midline. Black marking at caudal-
fin base (sometimes faint) made of a round or
vertically elongated black spot at extremity of
lateral line, a spot at base of lowermost principal
rays and an elongated, oblique blotch along base
of upper rudimentary rays. Two or three bars
across top of head, anterior one across eye and
continuing on cheek. Snout dark brown.
All fins hyaline. Dorsal fin with a large dark
grey blotch at base of anterior rays and two or
three irregular rows of spots on rays. Caudal fin
with 3-5 irregular rows of spots on rays. In a few
of the largest specimens, pelvic and anal fins
sometimes with a faint row of spots, pectoral fin
with blackish 2-3 anterior rays.
In juveniles (about 20-30 mm SL), bars nar-
rower, as wide as interspaces, not connected at
level of lateral line; pattern at caudal-fin base
distinct.
Distribution. Nemacheilus papillos is presently
known only from the lower Batang Hari drainage
and the adjacent Sungai Lalang (or Lalan, depend-
ing of maps) in the Musi drainage.
Etymology. Papilla (Latin), bud; os, oris (Latin),
mouth. A noun in apposition.
Remarks. Nemacheilus papillos is distinguished
from all species of Nemacheilus in having papil-
lated lips (vs. smooth or slightly plicated). It is
also distinguished from most in having the ante-
rior nostril at the tip of a short tube (vs. in the
anterior side of a small flap-like tube); this last
character is shared with N. selangoricus Duncker,
1904, N. spiniferus Kottelat, 1984, N. longipectoralis
Popta, 1905, N. tuberigum Hadiaty & Siebert, 2001
and N. pfeifferae (Bleeker, 1853a). The pattern of
black marks at the base of the caudal fin is shared
only with N. saravacensis Boulenger, 1894, from
which it is distinguished by having 9+8 branched
caudal-fin rays (vs. 8+8) and the colour pattern
of 10-11 bars connected at the level of the lateral
line (vs. 13-17 irregular blotches along lateral line,
more or less connected with saddles along the
dorsal midline; see Kottelat et al., 1993: pl. 26).
Nemacheilus papillos also has similarities with
N. selangoricus and N. spiniferus (elongated upper
caudal lobe, colour pattern on caudal and dorsal
fins, black bar across eye), but it is distinguished
in missing the acuminated scales on the caudal
peduncle, in missing the suborbital flap in male,
and in having a black pattern at caudal-fin base
made of an upper, a median and a lower spots
(vs. lower and median spots continuous).
Nemacheilus papillos is also distinguished from
N. tuberigum in missing the acuminated scales on
the caudal peduncle and in missing the suborbital
flap in male (data for N. tuberigum from Hadiaty
& Siebert, 2001).
Modigliania papillosa Perugia, 1893 is a species
of the genus Nemacheilus whose identity is still
unclear. It is distinguished from N. papillos by the
presence of a suborbital flap (vs. absence) (Kot-
telat, 1984).
Nemacheilus jaklesii was originally described
from Payakumbuh (Indragiri drainage, Sumatra).
As described and figured by Bleeker (1852c: 604;
1863-64: 7, pl. 103 fig. 5), it has a colour pattern
(13-20 regular bars on body) and the black basal
caudal marks somewhat similar to that of N. fas-
ciatus.
See Kottelat (1984) for comparison material.
Besides, we have examined 23 additional samples
of Nemacheilus from throughout Sumatra and have
not observed any with papillated lips. The exam-
ined material includes topotypes of N. pfeifferae
and N. papillosa and localities in the area of the
type locality of N. jaklesii.
Nemacheilus pfeifferae
Remarks. Highland species only known from
Sumatra (Kottelat et al., 1993). On caudal pedun-
cle, on each side of lateral line, a row of ovoid
Tan & Kottelat: Fishes of Batang Hari
57
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
scales, each with a tubercle at tip of exposed part;
a less developed form of the acuminate scales of
N. selangoricus and N. spiniferus but similar to the
condition described in N. tuberigum by Hadiaty
& Siebert (2001). On the flank, between the lat-
eral line and the base of the pelvic fin, there are
several rows of scales in which each scale has a
small tubercle.
Family Botiidae
Syncrossus reversus
Remarks. Bleekers figure of S. hymenophysa in
the Atlas (1863-64: pl. 102 fig. 3) shows a fish with
10 branched rays and no blotch at tip of dorsal
fin, which actually is S. reversus. The original
description of S. hymenophysa (Bleeker, 1852c) is
based on a single specimen from Palembang
(which is in the lowlands) with 12 or 13 branched
dorsal-fin ray and thus is not S. reversus, which
has 10-11 branched rays and inhabits the uplands
areas.
Family Cobitidae
Pangio atactos, new species
(Fig. 42)
Holotype. MZB 10995, 41.2 mm SL; Sumatra:
Jambi: Danau Kamining near Kampung Trasos,
ca. 5 km to southwards on unpaved road turning-
off from Muara Bungo – Muara Tebo road at km
36; M. Kottelat & H. H. Tan, 31 May 1994.
Paratypes. MZB 10996, 2, 22.1-33.0 mm SL; ZRC
51763, 5, 23.3-44.5 mm SL; CMK 11132, 5, 18.9-
45.6 mm SL; same data as holotype. – ZRC 51764,
2, 48.6-50.6 mm SL; Sumatra: Jambi: Sungai Pijoan,
upriver of Pijoan; from aquarium fish collectors;
M. Kottelat, 8 June 1994. – ZRC 42421, 2, 17.5-
44.5 mm SL; second stream on road towards
Muara Bulian, near junction at crossroads towards
Palembang, Muara Bulian and Jambi (1°47'44.4"S
103°25'16.1"E); H. H. Tan et al., 7 June 1996.
Diagnosis. Pangio atactos is distinguished from
all other congeners by its distinctive body colour
pattern consisting of 7-9 irregular triangular sad-
dles along the dorsal midline alternating with a
row of spots or blotches on the middle or lower
half of the body. Each saddle usually has a pale
rounded or transverse central area on dorsal
midline. It has similarity only with P. superba in
which the saddles are divided by a longitudinal
pale area forming a more or less continuous pale
stripe from the head to the base of the caudal
fin.
Description. General body shape and appearance
are shown in Figure 42. All large size specimens
are strongly arched, so that measurements are
difficult to duplicate. Morphometric data of
holotype (41.2 mm SL) are given here, only with
indicative value: total length 47.2 mm; head length
6.4 mm (15.5 % SL); predorsal length 26.1 mm
(63.3 % SL); prepelvic length 22.4 mm (54.4 % SL);
preanal length 31.8 mm (77.2 % SL); body depth
at dorsal origin 4.0 mm (9.7 % SL); depth of cau-
dal peduncle 3.6 mm (8.7 % SL); length of caudal
peduncle 8.8 mm (21.4 % SL) (2.4 times its depth);
body width 2.5 mm (6.1 % SL); length of pelvic
fin 4.0 mm (9.7 % SL); length of pectoral fin
4.6 mm (11.2 % SL); snout length 2.3 mm (5.6 %
SL, 36 % HL); eye diameter 0.9 mm (2.2 % SL,
14 % HL).
Dorsal fin inserted between pelvic and anal
fins, with 2 rudimentary, 1 simple and 6 branched
rays (and sometimes an additional simple ray);
first dorsal pterygiophore inserted behind neural
spine of vertebrae 28-30 (mode 29). Pectoral fin
with 8 rays. In males, pectoral fin curled upwards,
second ray 2-3 times thicker than other rays.
Pelvic fin with 7 rays, origin of first ray below
vertebral centra 25-27 (mode 26). Anal fin in-
serted shortly behind behind vent, with 2 rudi-
mentary, 1 simple and 5 or 5 {
branched rays;
first pterygiophore inserted behind hemal spine
of vertebrae 34-36 (mode 35). Caudal truncate or
emarginate, with 8+8 principal rays (7+7
branched).
Body entirely scaled, except belly between
pectoral fins. Head naked. Suborbital spine bifid,
outer branch straight, inner branch slightly
curved, longer and stronger than outer one. Three
pairs of barbels, one rostral, one maxillary and
one at each angle of mouth, reaching to eye.
Lower lip interrupted medially, each half with
an inner thickened lobe, not ending in a barbel-
like pointed tip; margin of membrane connecting
this lobe and barbel at corner of mouth entire.
Anterior nostril at tip of a short conical tube, not
barbel-like.
Vertebrae: 34 + 15 = 49 (1), 34 + 16 = 50 (1),
35 + 14 = 49 (2), 35 + 15 = 50 (3), 36 + 15 = 51 (1).
58
Fig. 42. Pangio atactos, MZB 10995, holotype, 41.2 mm SL; Sumatra: Batang Hari drainage, Danau Kamining.
Vertebrae 5 to 25-27 (mode 26) with pleural
ribs.
Coloration. Body yellow to orange in life, with
black markings. Three bars on head, first imme-
diately in front of eye, second immediately behind
eye, connected by a black area around eye; third
bar through posterior part of opercle. Sometimes
a black spot at tip of snout, not obvious. 7-9 sad-
dles along back, irregularly shaped, continued
ventrally by a narrower vertical bar (or triangular)
extending into lower half of body but not reach-
ing ventral midline. Usually a pale rounded or
transverse central area on dorsal midline in each
saddle. A row of spots or blotches on lower half
of flank (occasionally some on middle of flank),
irregularly shaped, positioned alternating with
saddles on back, opposite to spaces between sad-
dles, reaching ventral midline on caudal pedun-
cle. Last bar forming a large black blotch at
posterior extremity of caudal peduncle and at
base of caudal fin . A broad black bar about in
middle of caudal fin, in most largest specimens
branched at both extremities. A row of black spots
at midlength of dorsal-fin rays and membranes;
black saddles along dorsal-fin base extending
onto fin. Dark pigments in distal areas of rays
and membranes of anal, pelvic and pectoral
fins.
Juveniles (below about 27 mm SL) with more
contrasted pattern; bars thinner, saddles triangu-
lar, without the median pale area, and spots in
lower half of body small or absent.
Distribution. Pangio atactos is presently known
from three localities only in the Batang Hari drain-
age.
Etymology. From the ancient Greek ataktos,
meaning undisciplined, irregular, disorderly, not
in battle-order; allusion to the irregular disposi-
tion of the body markings. An adjective.
Remarks. With its irregular barred pattern on
the body and the black bar in the middle of the
caudal fin, P. atactos belongs to the P. shelfordii
(Popta, 1903) group of Kottelat & Lim (1993). In
this group, only P. superba (Roberts, 1989) has
saddles with an elongate projections below the
lateral midline and a row of dark marks in the
lower part of the body (not always present). Large
specimens of P. atactos and P. superba are distin-
guished by the number and shape of saddles
(7-9, triangular, vs. 6-7, longitudinally elongated
in lateral view; count does not include the ex-
panded third bar of head, located on the opercle
and nape). The saddles are well individualised
in P. atactos while they are often almost in contact
in P. superba. On the dorsal midline, each saddle
has a pale rounded or transverse central area in
P. atactos while in P. superba they are divided by
a longitudinal pale area forming a more or less
continuous pale stripe from the head to the base
of the caudal fin. The marks on the lower half of
the body are distinct along the whole body in
P. atactos, each being a distinct blotch alternating
with the saddles; in P. superba, these marks are
Tan & Kottelat: Fishes of Batang Hari
59
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 43. Pangio bitaimac, MZB 10997, holotype, 81.8 mm SL; Sumatra: Batang Hari drainage, Sungai Alai (right
side, reversed).
mostly behind the pelvic fins and do not have a
distinct shape and position.
As recognised by Kottelat & Lim (1993),
P. shelfordii in fact is an assemblage of several
species; this will be discussed elsewhere. Anyway,
none of them has an appearence similar to that
of P. atactos and they are not discussed here.
Pangio alternans Kottelat & Lim, 1993, from
the Mahakam drainage in Borneo also has a pat-
tern of saddles alternating with blotches in the
lower part of the body, but it has a much more
compact appearance, the saddles are more ir-
regularly shaped, the caudal fin is hyaline (vs.
with a black bar in the middle in P. atactos), the
tip of the snout has an obvious dark blotch, there
is a single broad bar through eye and it has
fewer vertebrae (32-34 + 13-15 = 45-48 vs. 34-36
+ 14-16 = 49-51).
Comparison material. Pangio superba. CMK 11584, 6;
ZRC 38787, 3, 30.8-44.1; Borneo: Kalimantan Barat:
Kapuas drainage, Sungai Letang near near Jongkong.
See Kottelat & Lim (1993) for additional material.
Pangio bitaimac, new species
(Fig. 43)
Holotype. MZB 10997, 81.8 mm SL, female; Su-
matra: Jambi: Sungai Alai; from aquarium fish
collectors, 1 Oct 2007.
Paratypes. MZB 10998, 15; ZRC 51765, 22; CMK
20840, 15; 62.6-95.4 mm SL; same data as holotype.
– ZRC 29161, 7, 52.0-80.4 mm SL; Sumatra: Jambi;
from aquarium fish collectors; 10 Nov 1992. – ZRC
42036, 16; CMK 17813, 6; 49.7-79.6; Sumatra:
Jambi: Sungai Alai; from aquarium fish collectors;
H. H. Tan et al., July 1997. – ZRC 43230, 1, 77.3 mm
SL; Sumatra: Jambi: Sungai Alai; from aquarium
fish collectors; T. Sim, July 1998. – CMK 11189, 3,
34.8-68.7 mm SL; Sumatra: Jambi: Sungai Alai,
km 28 on road from Muara Bungo to Muara Tebo;
from aquarium fish collector, 2 June 1994.
Additional material (non types). ZRC 38237, 1, 84.9 mm
SL; Malaysia: Johor: Kota Tinggi, Sungai Mupor; H. H.
Tan et al., 21 Aug 1994.
Diagnosis. Pangio bitaimac belongs to the P. an-
guillaris group of Kottelat & Lim (1993), defined
by a very elongate, worm-shaped body (body
depth 14-21 times in SL) and 46-52 + 15-20 = 61-71
vertebrae. It is distinguished from the other spe-
cies of this group by having the origin of the anal
fin behind the base of the dorsal fin (vs. below).
Further, it has 7+7 branched caudal-fin rays (vs.
6+6 in P. doriae), absence of scales on the cheek
(vs. present in P. doriae), the tube of the anterior
nostril developed into a nasal barbel (vs. not
developed into a nasal barbel in P. anguillaris).
Description. General body shape and appearance
are shown in Figure 43. All large size specimens
are strongly arched, so that measurements are
difficult to duplicate. Morphometric data of
holotype (81.8 mm SL) are given here, only with
indicative value: total length 86.9 mm (106.2 %
SL); head length 7.9 mm (9.7 % SL); predorsal
length 62.0 mm (75.8 % SL); prepelvic length
51.8 mm (63.3 % SL); preanal length 65.1 mm
(79.6 % SL); body depth at dorsal-fin origin
5.2 mm (6.4 % SL); length of caudal peduncle
14.4 mm (17.6 % SL); depth of caudal peduncle
3.3 mm (4.0 % SL) (4.4 times in its length); body
width 3.9 mm (4.8 % SL); length of pelvic fin
1.8 mm (2.2 % SL); length of pectoral fin 2.3 mm
(2.8 % SL); snout length 3.1 mm (3.8 % SL, 39 %
60
HL); eye diameter 0.6 mm (0.7 % SL, 7 % HL).
Dorsal fin with 2 rudimentary, 1 simple, 6
branched and 1 unbranched rays, last 2 articulat-
ing on same pterygiophore; first pterygiophore
inserted behind neural spine of vertebrae 43-45
(mode 44). Pectoral fin with 6-7 rays. In males,
pectoral fin curled upwards, second ray about
3-4 times thicker than other rays. Pelvic fin with
5 rays, origin of first ray below vertebral centra
38-39. Anal fin inserted almost immediately be-
hind vent, origin behind base of dorsal fin, with
2 rudimentary, 1 simple, 5 branched and 1 un-
branched rays, last two articulating on same
pterygiophore; first pterygiophore inserted be-
hind hemal spine of vertebrae 46-48 (mode 47).
Caudal truncate to very slightly emarginate, with
8+8 principal rays (7+7 branched).
Body entirely scaled, in some specimens naked
between pectoral fins. Head naked. Suborbital
spine bifid, outer branch straight, inner branch
slightly curved, longer and stronger than outer
one. Three pairs of barbels, one rostral, one max-
illary and one at angle of mouth, reaching slight-
hly behind eye. Lower lip interrupted medially,
each half with an inner slightly thickened lobe,
not ending in a barbel-like pointed tip; margin of
membrane connecting lobe and barbel at corner
of mouth entire, not fringed, without notches.
Anterior nostril at tip of a short conical tube,
posterior edge elongated into a nasal barbel, ex-
tending beyond eye.
Vertebrae: 46 + 16 = 62 (1), 47 + 14 = 61 (1),
47 + 15 = 62 (2), 47 + 16 = 63 (1), 48 + 15 = 63 (2),
48 + 16 = 64 (1). Kottelat & Lim (1993: 246) count-
ed 46-49 + 15-17 = 63-64 in 7 additional specimens.
Vertebrae 5 to 46-47 (mode 47) with pleural
ribs.
Coloration. Preserved specimens: Body yellow-
ish brown. Dorsal half of body and whole caudal
peduncle usually very finely dotted (each dot
being a dark greyish to black scale), denser on
middle of body and forming a vague midlateral
stripe. Stripe more conspicuous on caudal pedun-
cle and above pectoral fin. Rest of body usually
without pigments. Dorsal part of head plain yel-
lowish brown. Tip of snout, part of space between
nostrils, and between nostrils and mouth blackish.
Cheek dotted. A short black stripe on both sides
of groove of suborbital spine usually present.
Caudal fin with black pigments on unbranched
part of rays and on proximal area of all interra-
dial membranes, resulting in a vertically elon-
gated blotch. Black marks along dorsal-fin rays
in some specimens. Other fins hyaline.
In life: violet.
Distribution. The species is presently known
only from the Batang Hari drainage in Jambi
province, Sumatra. We have examined a single
specimen from near Kota Tinggi at the southern
extremity of the Malay Peninsula (ZRC 38237),
which is possibly conspecific. We have not col-
lected the species ourself and cannot comment
on the habitat. However, the violet live colour
and the very reduced black pimentation on the
flank, shared with a number of other fish species
that live burried in the mud (e.g. Caragobius
urolepis), suggest that P. bitaimac could have a
similar mode of life.
Etymology. The name bitaimac is the Latinized
form of bee tai mak, a kind of short and thick rice
noodle, commonly consumed locally in Southeast
Asia. A noun in apposition.
Remarks. Kottelat & Lim (1993: 246) recognised
4 species groups within Pangio, based on vertebrae
counts, body shape and colour pattern. They
recognised a P. anguillaris group including the
species with a large number of vertebrae (46-52
+ 15-17 = 61-71, vs. 31-40 + 12-16 = 45-55 in the
other groups). They included three species in this
groups: P. anguillaris, P. doriae and P. pulla. Pangio
pulla is very distinctive in having a banded black
and yellowish colour pattern (vs. plain brown)
and a deeper body (11-12 times in SL, vs. 14-21)
than the other species of the group. Its vertebrae
count (46-48 + 15-16 = 61-62) is at the lower limit
of the range observed in the group. We think that
it does not belong to this group but cannot place
it in any of the three other groups and it is prob-
ably better considered a group of its own. Pangio
pulla is also distinguished by the absence of pel-
vic fins, but this character appears in different
lineages of Pangio and can only be used for the
diagnosis of some species, not lineages.
Material of P. bitaimac was identified as P. do-
riae (Perugia, 1892) by Kottelat & Lim (1993), but
they already pointed to the difference in the
number of branched caudal-fin rays between them
and the remaining material of P. doriae (7+7, vs.
6+6). The availability of more material shows that
these indeed are distinct species.
Besides the number of caudal-fin rays, P. bitai-
mac is distinguished from P. doriae by the colour
Tan & Kottelat: Fishes of Batang Hari
61
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
pattern at the base of the caudal fin (black pig-
ments on proximal area of all interradial mem-
branes, resulting in a vertically elongated blotch,
vs. black pigments restricted to the interradial
membranes between the 4 median rays, extending
on most of membrane, resulting in a short stripe
continuing the stripe on caudal peduncle), the
shape of the caudal fin (truncate or very slightly
emarginate, vs. distinctly emarginate), the origin
of the anal fin clearly behind the base of the last
dorsal-fin ray (vs. in front or below) and rela-
tively shorter nasal barbel (the tube part of the
anterior nostril is about | of the total length of
the nasal barbel, vs. about ƒ-~).
Pangio bitaimac differs from P. anguillaris (Vail-
lant, 1902) in having the anterior nostril modified
into a nasal barbel (vs. at tip of a short tube);
fewer vertebrae (46-49 + 14-17 = 61-67, vs. 50-
52 + 19-20 = 69-71; Kottelat & Lim, 1993: 246) and
related to a shorter distance between pelvic and
anal fins (14.9-18.4 % SL, vs. 18.6-22.0; measured
on 10 P. bitaimac, ZRC 51765 and 12 P. anguillaris,
ZRC 38629, 38586, 34884, from Sumatra and Pa-
hang), the origin of the anal fin behind the base
of the last dorsal-fin ray (vs. in front), and by the
colour pattern at the base of the caudal fin (black
pigments on proximal area of all interradial
membranes, resulting in a vertically elongated
blotch, vs. black pigments restricted to the inter-
radial membranes between the 4 median rays,
extending on most of membrane, resulting in a
short stripe continuing the stripe on caudal pe-
duncle). Kottelat & Lim (1993: 214) erroneously
report 6+6 branched caudal rays in P. anguillaris;
we re-examined their material and observed 7+7
in all specimens.
Roberts (1989) and Kottelat & Lim (1993)
considered Cobitophis perakensis Herre, 1940 (de-
scribed from Perak, Malay Peninsula) as a syno-
nym of P. anguillaris, based on the examination
of the types and new material from the Malay
Peninsula and Borneo. They also list Acanthoph-
thalmus vermicularis Weber & de Beaufort, 1916
(described on the basis of a single specimen from
the Kampar Kiri in Sumatra) as a synonym. This
is the only record of P. anguillaris from Sumatra
prior to our surveys. Weber & de Beaufort (1916:
34) record the presence of only 3 pairs of barbels
(that is, no nasal barbels) and this excludes the
possibility that P. bitaimac is identical with A. ver-
micularis (that is, P. anguillaris). We have not seen
differences between our P. anguillaris from Bor-
neo, Sumatra and the Malay Peninsula and we
retain P. vermicularis and P. perakensis in the syn-
onymy of P. anguillaris.
The P. anguillaris illustrated in Kottelat et al.
(1993: pl. 28) and Kottelat & Lim (1993: 214) is a
specimen from Nan (Chao Phraya drainage,
Thailand). It is not certain that it is conspecific
with the Sundaic populations.
Pangio lumbriciformis Britz & Maclaine, 2007
possibly also belongs to the P. anguillaris group.
It shares the dark grey to black stripe on median
caudal-fin rays. It differs from P. bitaimac in hav-
ing fewer vertebrae (40-41 + 13-15 = 54-56, vs.
46-49 + 14-17 = 61-67).
Comparison material. Pangio anguillaris: ZRC 38629,
5, 56.3-65.1 mm SL; CMK 11187, 11, 56.7-62.0 mm SL;
Sumatra: Jambi: Sungai Alai, near Muara Bungo. – ZRC
38586, 2, 40.2-47.2 mm SL; Sumatra: Jambi: Danau Ka-
mining near Muara Bungo. – ZRC 34884, 5, 57.3-66.2 mm
SL; Malaysia: Pahang: Sungai Salan at Jerantut. – ZRC
1442, 3 paratypes of C. perakensis, 50.3-54.7 mm SL;
Malaysia: Perak: Chenderoh dam.
Pangio cf. bitaimac: ZRC 38237, 1, 86.3 mm SL;
Malaysia: Johor: Kota Tinggi, Sungai Mupor.
Pangio doriae: ZRC 38793, 4, 40.2-55.9 mm SL; CMK
11632, 9, 40.4-61.1 mm SL; Borneo: Kalimantan Barat:
Kapuas drainage: Sungai Embau.
Pangio semicincta
Remarks. Kottelat & Lim (1993: 229-230) tenta-
tively considered P. kuhlii (Valenciennes, in Cu-
vier & Valenciennes, 1846) and P. semicincta as
synonyms. Fresh material from Java shows that
the two species have very distinctive colour pat-
terns and Kottelat (1995b; Kottelat & Whitten,
1996a) recognised P. kuhlii endemic to Java and
P. semicincta from Sumatra, Borneo and the Malay
Peninsula. Fraser-Brunner (1940) described P. su-
matrana on the basis of a single specimen from
Lahat in the Musi drainage (Sumatera Barat
Province). It has been tentatively placed in the
synonymy of P. kuhlii by Kottelat & Lim (1993).
We have now obtained material of Pangio from
additional localities in the Musi drainage, includ-
ing from the vicinity of Lahat (type locality of
P. sumatrana). We did not observe differences
between them and the populations of P. semi-
cincta from elsewhere in Sumatra; therefore we
consider them as synonyms. As they are simul-
taneous synonyms, we give precedence to P. semi-
cincta.
Fraser-Brunner distinguished his P. kuhlii
sumatrana from P. k. kuhlii on the basis of colour
62
pattern. The character that he mentioned (body
with about 12 broad bands, narrowing somewhat
below) corresponds to some of our Musi speci-
mens. Besides, he distinguished P. kuhlii (includ-
ing his P. k. sumatrana) from P. semicincta by
having a smaller head (8 times in length, vs. 6.5),
the pelvic fin well behind the middle of the
length (vs. in middle of length of fish, including
caudal fin), the dorsal fin with 7-8 branched rays
(vs. 6), and the anal-fin origin slightly behind the
end of the dorsal fin (vs. immediately below end
of dorsal fin). This is contradicted by his drawings,
which do not show difference in fin positions
between his P. semicincta and P. kuhlii sumatrana.
Our specimens from near the type locality of
P. sumatrana have 6 { branched dorsal-fin rays as
does P. semicincta, and the head length is 6.1-7.0
times in total length.
Material. ZRC 51766, 5; CMK 20538, 5; 29.0-48.0 mm
SL; Sumatra: Sumatera Selatan: Musi drainage: Kikim
River at Bunga Mas, about 40 km west of Lahat.
Family Osphronemidae
Trichopodus leerii
Trichopodus trichopterus
Remarks. These two species have usually been
placed in Trichogaster. When originally created,
Trichogaster Bloch (in Schneider, 1801: 164) in-
cluded two nominal species, of which Trichogaster
fasciatus Bloch (in Schneider, 1801: 164) was sub-
sequently designated as type species by Jordan
& Evermann (1917: 58). Trichogaster fasciata is a
member of the genus usually called Colisa Cu-
vier (in Cuvier & Valenciennes, 1831: 359), and
this makes Trichogaster the valid name of this
genus. Contrary to comments by Derijst (1997:
222), Colisa is not available from Hamilton (1822)
as this name is clearly not intended as a scien-
tific name (Code art. 1.1 and Glossary).
The type species of Trichopodus La Cepède
(1801: 125) is Labrus trichopterus Pallas (1770: 45),
by subsequent designation by Bleeker (1879: 21)
Contrary to statement in Derijst (1997: 223), Cu-
vier (in Cuvier & Valenciennes, 1831: 388) did not
designate a type species for Trichopodus. He used
the wording le véritable trichopode and this
does not constitute a type species designation.
Family Tetraodontidae
Monotrete leiurus
Remarks. Material usually identified as M. lei-
urus exhibits a great variability and several
nominal species earlier treated as synonyms have
recently been revalidated and additional species
have been described (Kottelat, 1995a, 2000; Rob-
erts, 1998). It is still not clear how many species
occur in Java, Sumatra and Borneo. Monotrete
leiurus was described on the basis of 5 specimens
from Batavia (now Jakarta) that have subse-
quently been mixed with other samples. The
potential syntypes examined by Dekkers (1975)
are in poor condition and have lost their colour
pattern (a major diagnostic character within this
group).
Bleeker (1851b: 97) described the type series
from Batavia (Jakarta), in sea and estuaries. The
species now identified as M. leiurus inhabits
freshwater and has never since been recorded in
estuaries or in the sea.
We examined recent specimens from western
Java (MZB 1330, 1332; Fig. 44). They have a large
round dark brown blotch on the flank, with a
round reddish spot in the center, under the
dorsal-fin origin, surrounded by a ring of some-
what smaller blotches, and the rest of the body is
covered by similar, smaller, blotches. In specimens
about 35 mm SL the blotches forming the outer
ring are very close and in the smallest examined
specimen (30 mm SL) they form a more or less
continuous circle. There is usually a pale brown,
unspotted patch in the upper part of the caudal
peduncle and a broad pale brown band along the
upper edge of the caudal fin. There is no marking
on the belly. The snout is long, about twice the
diameter of the eye. We observed the character
states in material from Sumatra (Musi and Batang
Hari drainages, Lampung) and the Malay Penin-
sula as far north as at least Pahang. The red spot
of the ocellus becomes darker and less distinct
with increasing size. It is missing in the 84.8 mm
SL Batang Hari and the 76.0 mm SL Musi speci-
mens and in all specimens above 60 mm SL in
the Bogor material (MZB 1330); above this size
the blotch is totally black. But it is still present in
the 102.6 mm SL Pahang specimen, although very
dark.
For the time being we consider these popula-
tion groups (Java, Sumatra, Malay Peninsula) as
conspecific and call them M. leiurus. This identi-
Tan & Kottelat: Fishes of Batang Hari
63
Ichthyol. Explor. Freshwaters, Vol. 20, No. 1
Fig. 44. Monotrete leiurus, MZB 1332, 54.1 mm SL; Java: Bogor (right side, reversed).
fication may have to be revised when more mate-
rial (larger series) become available. On Borneo,
the few samples we have seen from the Barito
drainage are possibly conspecific, although they
seem to have a well marked hump in the middle
of the back. They differ in having the patterning
of the blotches less regular in distribution (the
ring about the subdorsal blotch is not as con-
spicuous as in the Javanese specimens), the
blotches are less regularly shaped, in some
specimens almost polygonal, with narrower
spaces between the blotches. At least part of the
material earlier reported as Tetraodon leiurus from
the Mahakam drainage (eastern Borneo) is a
distinct species, M. hilgendorfii (Popta, 1905; see
Kottelat, 1995a). It is distinguished by the longi-
tudinally elongated blotches on the belly and
lower part of the flank; there is no ocellus and
there is no conspicuous large black blotch under
the origin of the dorsal fin.
In our material from the Kapuas drainage
(western Borneo), small specimens (less than